Skip to main content

Full text of "Flora of Guatemala"

See other formats


RARY 
OF THE 

UNIVERSITY 
OF ILLINOIS 

580-5 



BIOLOGY 



Return this book on or before the 
Latest Date stamped below. 

Theft, mutilation, and underlining of books 
arc reasons for disciplinary action and may 
result in dismissal from the University. 
University of Illinois Library 



FEb 9 1)65 



L161 O-1096 



FLORA OF GUATEMALA 



PAUL C. STANDLEY 



AND 



JULIAN A. STEYERMARK 



FIELDIANA: BOTANY 
VOLUME 24, PART I 

Published by 

CHICAGO NATURAL HISTORY MUSEUM 
AUGUST 29, 1958 

SEP 



FLORA OF GUATEMALA 



FLORA OF GUATEMALA 



PAUL C. STANDLEY 

Curator Emeritus of the Herbarium 



AND 



JULIAN A. STEYERMARK 

Formerly Curator of the Herbarium 



FIELDIANA: BOTANY 

VOLUME 24, PART I 

Published by 

CHICAGO NATURAL HISTORY MUSEUM 
AUGUST 29, 1958 



Published with the assistance of a grant from 

THE LILLY RESEARCH LABORATORIES, ELI LILLY AND COMPANY 

INDIANAPOLIS, INDIANA 



Library of Congress Catalog Card Number: ^8-8076 



PRINTED IN THE UNITED STATES OF AMERICA 
BY CHICAGO NATURAL HISTORY MUSEUM PRESS 



CONTENTS 



Families Included in Part I 



Cycadaceae ..................... 11 

Podocarpaceae .................. 20 

Araucariaceae ................... 23 

Cupressaceae .................... 26 

Pinaceae ....................... 36 

Taxodiaceae .................... 56 

Taxaceae ....................... 60 

Typhaceae ...................... 63 

Pandanaceae .................... 67 

Potamogetonaceae ............... 68 

Najadaceae ..................... 73 

Alismataceae . 75 



Butomaceae 

Hydrocharitaceae 

Triuridaceae 

Cyperaceae 

Palmae 

Cyclanthaceae 

Araceae ....... 

Lemnaceae 
Mayacaceae 
Xyridaceae 
Eriocaulaceae 
Bromeliaceae . . 



PAGE 

80 

84 

86 

90 

196 

299 

304 

364 

369 

370 

374 

380 



LIST OF ILLUSTRATIONS 
TEXT FIGURES 

PAGE 

1. Ceratozamia mexicana .... 13 

2. Zamia Loddigesii 17 

3. Zamia Tuerckheimii 20 

4. Podocarpus oleifolius . . 24 

5. Cupressus lusitanica . . 29 

6. Juniperus Standleyi . . .35 

7. Abies guatemalensis 39 

8. Pinus Ayacahuite 45 

9. Pinus caribaea .... . . 47 

10. Pinus Montezumae ... 49 

11. Pinus oocarpa . . 52 

12. Pinus pseudostrobus .54 

13. Pinus Strobus var. chiapensis . .56 

14. Taxodium mucronatum 59 

15. Taxus globosa 62 

16. Typha domingensis .... .65 

17. Echinodorus grandiflorus 77 

18. Sagittaria lancifolia .81 

19. Hydrocleis Standleyi . 83 

20. Vallisneria americana .87 

21. Sciaphila picta . . 89 

22. Bulbostylis capillaris .94 

23. Calyptrocarya glomerulata . 99 

24. Carex polystachya 105 

25. Cyperus rotundus . . . . 137 

26. Dichromena Watsoni . . . 144 

27. Eleocharis elegans .... .... 148 

28. Fimbristylis dichotoma .... .... 158 

29. Hypolytrum Schraderianum . 164 

30. Kyllinga pumila . 167 

31. Rhynchospora tenuis ... .177 

32. Scirpus cubensis var. gracilis . . . 182 

33. Scleria pterota . 191 

34. Uncinia ham at a .195 

35. Acrocomia mexicana . 203 

36. Asterogyne Martiana 205 

37. Astrocaryum mexicanum ... 207 

38. Bactris balanoidea .... .210 

39. Bactris trichophylla . 212 

vii 



PAGE 

40. Calyptrogyne Donnell-Smithii 216 

41. Chamaedoreaelegans, C. Ernesti-Augustii, C. Tepejilote, C. Arenbergiana, 

C. Rojasiana, C. geonomaeformis, C. oblongata, C. Pachecoana . 226, 227 

42. Cryosophila argentea 257 

43. Desmoncus ferox ... 261 

44. Geonoma Seleri 270 

45. Manicaria saccifera . . 272 

46. Orbignya cohune ... 275 

47. Paurotis Wrightii . . ... 278 

48. Reinhardtia gracilis ... 282 

49. Sabal mexicana ... . . 288 

50. Scheelea Preussii .... ... 292 

51. Synechanthus fibrosus . . ^ 296 

52. Thrinax parviflora . . . 297 

53. Carludovica utilis .... 303 

54. Anthurium crassinervium 311 

55. Dieffenbachia seguina ... 327 

56. Monstera pertusa 333 

57. Montrichardia arborescens , 336 

58. Philodendron Hoffmannii 343 

59. Pistia Stratiotes 348 

60. Spathipyllum blandum 351 

61. Xanthosoma robustum 361 

62. Lemna paucicostata, L. p. var. membranacea, L. valdiviana, 

Wolffia Welwitschii 367 

63. Mayaca fluviatilis 370 

64. Xyris Jupicai 373 

65. Aechmea magdalenae 385 

66. Aechmea tillandsioides var. Kienastii 386 

67. Ananas comosus 388 

68. Androlepis Skinneri ... 390 

69. Billbergia viridiflora . . 392 

70. Bromelia Pinguin, B. sylvestris, B. Karatas 394 

71. Catopsis apicroides . . 397 

72. Catopsis cucullata . . 399 

73. Catopsis floribunda ... .... 400 

74. Catopsis Lundelliana . .... 402 

75. Catopsis Morreniana 403 

76. Catopsis nitida . .... 404 

77. Catopsis nutans . ... 405 

78. Catopsis sessiliflora . . .... 407 

79. Catopsis Wawranea . ... 409 

80. Greigia Steyermarkii .. 410 

81. Guzmania nicaraguensis 412 

82. Guzmania Scherzeriana var. guatemalensis 412 

83. Hechtia guatemalensis 414 

84. Hohenbergia guatemalensis . 416 

85. Lindmania micrantha .. 418 

86. Pitcairnia flagellaris ... 421 

87. Pitcairnia heterophylla . . . . 423 



PAGB 

88. Pitcairnia imbricata ... . 423 

89. Pitcairnia macrochlamys . 425 

90. Pitcairnia punicea ... 426 

91. Pitcairnia Tuerckheimii . 428 

92. Tillandsia anceps . 435 

93. Tillandsia argentea 435 

94. Tillandsia Baileyi . .437 

95. Tillandsia Balbisiana . . 438 

96. Tillandsia brachycaulos . 438 

97. Tillandsia bulbosa . . 439 

98. Tillandsia Butzii 440 

99. Tillandsia caput-Medusae . 442 

100. Tillandsia chlorophylla .442 

101. Tillandsia elongata var. subimbricata 445 

102. Tillandsia fasciculata . .446 

103. Tillandsia festucoides . . .447 

104. Tillandsia grandis . . . 449 

105. Tillandsia ionantha ... . 451 

106. Tillandsia juncea ... .452 

107. Tillandsia lampropoda . 453 

108. Tillandsia monadelpha . 455 

109. Tillandsia multicaulis . 456 

110. Tillandsia polita ... .457 

111. Tillandsia ponderosa 459 

112. Tillandsia pruinosa . 460 

113. Tillandsia punctulata . . 461 

114. Tillandsia recurvata .... 462 

115. Tillandsia Rothschuhiana . 463 

116. Tillandsia Schiedeana 464 

117. Tillandsia Standleyi . 466 

118. Tillandsia usneoides . 468 

119. Vriesia heliconioides . 472 

120. Vriesia montana ... 473 

121. Vriesia pectinata . 475 



IX 



Flora of Guatemala 



PLAN OF THE FLORA 

This Flora of Guatemala is intended to list all phanerogamic plants 
growing naturally in Guatemala and British Honduras. The latter 
country has been included, since on both geographic and botanical 
grounds it is essentially part of Guatemala. Because British Hon- 
duras was covered by a rather recent publication, Forests and flora of 
British Honduras by Samuel J. Record and Paul C. Standley (1936), 
its plant geography is not discussed in the present Flora, and the 
authors have not collected there. 

Besides the native and naturalized plants, there have been in- 
cluded in this Flora the common cultivated plants of Guatemala, 
whether of economic or ornamental value. These plants are mostly 
the same ones to be observed in cultivation in all other Central 
American countries and in tropical America generally. Such plants 
are more important to Guatemalan people than species of forests 
and meadows, and since there is no generally available publication 
treating the cultivated plants of tropical America, the information 
given here should be of service to them. Moreover, intelligent and 
curious strangers visiting the country, especially those without pre- 
vious tropical experience, are likely to be just as much interested in 
cultivated plants as wild ones, and will desire information regarding 
them. The agriculture of Central America and especially of Guate- 
mala is a subject well deserving detailed study. Some attention has 
been given in recent years to such crops as corn and beans, but a 
great deal of time could be spent profitably in investigating other 
economic plants, both cultivated and wild ones. 

All the work of preparing the Flora has been done at Chicago 
Natural History Museum, and is based primarily upon recent col- 



2 FIELDIANA: BOTANY, VOLUME 24 

lections obtained by four botanical expeditions dispatched to Guate- 
mala by the Museum. These have assembled some 54,000 collections 
of plants from almost every part of Guatemala. Besides our own 
collections, we have studied series of the basic and historical ones 
distributed by Captain John Donnell Smith, whose collections com- 
prised a large percentage of the many new species described from 
Guatemala. We have used also a set of the recent but highly im- 
portant collections made by Dr. A. F. Skutch, and have examined 
photographs or fragmentary specimens of types of many species 
described from the earlier collections made in Guatemala. Some of 
these are no longer extant, having been consumed by flames that 
destroyed the Berlin Herbarium in March of 1943. 

If one will examine the catalogue of collectors in this volume, it 
will be found that the amount of earlier botanical exploration in 
Guatemala, aside from the collections just mentioned, was singularly, 
small, especially when one considers the high reputation of the Gua- 
temalan flora. Few of the collections besides those mentioned were 
as large as 300 numbers. Most of them were made in selected areas, 
easy of access or else attractive because of special comfortable con- 
ditions existing there (often a finca owned by some immigrant from 
the United States or Germany), or because of special groups of plants 
that it was decided to study in some particular spot where they were 
known to occur. 

The collections made by Heyde and Lux and distributed by Cap- 
tain Smith were assembled at a wide range of localities, as were those 
of Dr. Eduard Seler and his wife, who were visiting archeological sites, 
and the series obtained by Dr. Skutch, who was interested primarily 
in birds. Karl T. Hartweg had the high privilege of being almost 
the first collector to visit Guatemala. He worked in several widely 
separated but easily accessible localities, but the plants he described, 
although highly important historically, were not numerous. The 
rest of the collections, at least those of collectors from the United 
States, were made principally along the railroad between Puerto 
Barrios and Guatemala, on Volcan de Agua, or where lodging was 
most comfortable on the coffee fincas of Alta Verapaz. 

The collections studied we believe to be adequate for preparation 
of a flora. We have borrowed from the United States National 
Museum, Gray Herbarium, and the University of Michigan a few 
types and critical specimens necessary to establish matters of tax- 
onomy, and to the curators who courteously lent this material we 
are greatly obliged. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 3 

It would seem that there are now reported most of the species 
that could be expected in Guatemala, practically all those found both 
north and south of that country, and the great majority of the 
Mexican species that could be expected to extend so far southward. 
However, work upon the Guatemalan flora has shown that plants 
of Mexico appear in Guatemalan localities where they are least 
expected, and it is not believed that their numbers have been ex- 
hausted. The borders of Guatemala have been fairly well explored, 
and for that reason intruding species from the north and south are 
less likely to be found. The Guatemalan flora as now compiled in- 
cludes almost every species known from El Salvador and Honduras, 
so that many additions from that direction can not be anticipated. 

There are certain parts of Guatemala where some additions can 
be found, without any doubt. Little is known of northern Pete'n, 
but in the parts of Mexico north of the Guatemalan border, them- 
selves imperfectly known, there are few known species that fail to 
reach Guatemala. The areas of Guatemala where additions are most 
likely to appear are northern Huehuetenango and Quiche^ where a 
good number of new species will be found, the mountains of Baja 
Verapaz, and the Cockscomb Mountains along the boundary between 
Guatemala and British Honduras. The southeastern portion of 
Jutiapa has not been explored, but it is improbable that many addi- 
tions will be found there. It is rather strange, but explainable, that 
the main portion of Baja Verapaz still remains unvisited or at least 
the plants have not been collected by a botanist. The most profit- 
able region remaining for botanical collecting we believe to be north- 
ern Quich^ and Huehuetenango, and some surprises are confidently 
to be expected there. We believe that the present enumeration con- 
tains the vast majority of plants native in Guatemala, and it will be 
a surprise if the number is increased through later exploration by 
10 per cent. 

The general form in which data of the systematic portion of the 
Flora are printed will be, as a rule, sufficiently obvious. Generally 
in the synonymy only the essential names are listed : the name-bring- 
ing synonym, if any, and synonymous names that have been used for 
Guatemalan plants or recently for those of Central America. There 
have been included all names known to have been based on Guate- 
malan material, and under each of these the type is cited. Free 
citation has been made also of synonymous names based on material 
from Central American countries other than Guatemala or from 
nearby portions of Mexico, especially when such names have not 
been referred previously to synonymy. An attempt has been made 



4 FIELDIANA: BOTANY, VOLUME 24 

to account for all names that have been reported from Guatemala, in 
either the synonymy or remarks after the formal description of the 
species. 

The names of authorities for species names have been abbreviated 
according to ordinary usage, except when the names are short. Au- 
thorities for the generic names are written in full, except in the case 
of "L." (Linnaeus or Linne") and "HBK." (Humboldt, Bonpland, 
and Kunth). The abbreviations for authors' names often mystify 
beginners in botanical work, and we have not supplied a list of their 
explanations, such as sometimes appears in a Flora. However, most 
of such abbreviated names will be found written in their full form 
after the name of a genus in some part of this Flora. 

Under each genus or family there have been cited the most use- . 
ful monographic accounts of the group, when such works exist. Some 
older monographs no longer of practical value have been omitted, 
although the synonymy of the species often will give a clue to them. 

We have given all accessible vernacular names used in Guatemala 
that we consider reasonably authentic. Others that have come to 
our attention have been omitted because of doubt regarding their 
form or application, often because of obviously erroneous transcrip- 
tion that it was impossible to correct. Erroneous names do more 
harm than good, and in general they should not be collected by per- 
sons without a fairly good knowledge of Spanish as it is spoken and 
written in America, not in textbooks. In gathering vernacular names 
they should be checked with two or more persons if at all possible. 
An even better method is to have them written down by the inform- 
ant, if he can write. His spelling may be bad, but at least he is likely 
to represent the sound of the word. Especially with Indian names 
it is easy for a person to misunderstand the sound of a word. Even 
educated Latin people sometimes are unable to decide whether the 
sound in a new word is, for example, "r" or "rr." 

We have collected most of the vernacular names. Others have 
been taken from labels of other collectors, and some from publica- 
tions whose accuracy we trust. A large number of names we have 
cited are from lists furnished by Don Jos Ignacio Aguilar, formerly 
Director of the Finca Nacional La Aurora, who has given much 
attention to the Guatemalan flora. We are indebted to Professor 
Ulises Rojas, formerly Director of the Jardin Botanico de Guate- 
mala, who has given help with vernacular names and economic data. 
The vernacular names entered here doubtless include some mistakes, 
even after the best of care, and this is particularly probable in case 
of non-Spanish terms. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 5 

There is much more to be done in the field of Guatemalan ver- 
nacular names. In published lists we have seen many plant names 
whose identity it is impossible to guess. The value of a particular 
vernacular name varies usually in proportion to the importance of 
the plant. If the plant is one of which some definite use is made, or 
a showy and conspicuous one, the vernacular name is likely to be 
fixed and in common usage. If the plant is inconspicuous, or if no 
use is made of it, the vernacular name often is open to suspicion and 
seldom is in common use. However, the local standards of impor- 
tance and individual ideas of beauty or conspicuousness may differ 
from those of the questioning botanical collector. It always is un- 
wise to press for a vernacular name, and it is much better to ask for 
one indirectly. Many people have a talent for manufacturing them 
on the spot and find great satisfaction in fooling a foreigner and boast- 
ing about it afterward. The senior author greatly admired the ability 
in this respect of a small boy at Jutiapa. He happened to have a 
remarkable knowledge of the plants of the region, but he was not 
content with what he really knew. He would invent a name almost 
as quickly as a new plant was found, and his names often were ex- 
traordinarily suitable and subtle too much so for belief. Adults in 
country districts think they will lose face if they can not supply a 
name for any plant found, and will often make a clumsy effort to 
manufacture one. And a great many Central Americans, like natives 
of the United States, make mistakes in recognition of plants, thus 
giving them incorrect names really belonging to different plants. 
People who recognize wild plants easily when in the ground, often 
are unable to place them when they see a detached branch in a work 
room. Woodsmen usually pay much more attention to the bark and 
trunk of a tree than to any other portion of it, and if reliable names 
for trees are desired, it always is better to obtain them from a quali- 
fied person standing by the tree in question. 

Besides vernacular names from Guatemala, we have cited those 
known from British Honduras as a matter of course, Maya names of 
Yucatan that may well be used by the people of Pete"n, and from 
Honduras, El Salvador, and the bordering states of Mexico many 
names that may be used in Guatemala. There have been cited occa- 
sionally herbarium names from more remote regions when these have 
not appeared previously in print. 

In describing Guatemalan ranges for species, there is cited each 
department from which it is definitely known. Most of the depart- 
ment records are based on herbarium specimens, a few on published 
records, and, in the case of well-known species, sometimes on field 



6 FIELDIANA: BOTANY, VOLUME 24 

notes, especially in the case of plants so common that specimens were 
not desired. The altitudinal distribution is given when known. Alti- 
tude is a dominant factor in distribution of the majority of the species, 
but for others, mostly weedy ones, it has little significance. In de- 
scribing the Guatemalan range of the species, the departments are 
cited in a definite order, beginning with Pete"n and proceeding to the 
North Coast, the Oriente, the central departments, northwestern 
Guatemala, and finally to the Occidente, ending with the Depart- 
ment of San Marcos. This order is approximate, since the depart- 
ments cannot be followed in a single line. 

In the descriptions of genera and species, words of Latin deriva- 
tion have been used intentionally, although doubtless not consist- 
ently. Latin terms obviously are more intelligible to Spanish-speaking 
people. In general, in writing the Flora two groups of people have 
been kept in mind : foreign botanists interested primarily in the iden- 
tity of the plants found in Guatemala, the means of distinguishing 
them, and data concerning their occurrence and uses within the 
country; and Guatemalans themselves, who may be interested in 
some other botanical phases, such as the relation of their plants to 
those of other countries. 

As in other Central American countries, the cryptogamic plants 
of Guatemala are still imperfectly known. A separate volume of this 
Flora contains an admirable account of Guatemalan mosses, a work 
unique in its field for the mainland of tropical North America, and 
perhaps also for South America. The lichens, hepatics, fungi, and 
algae are less known, and the only papers relating to them are of little 
use except to specialists in their groups. It is hoped that there may 
be included in the Flora an account of the ferns and their allies. 
These plants are usually collected and studied with the flowering 
plants, and treated in much the same manner. They are very numer- 
ous in Guatemala, and, of course, much better known than the lower 
cryptogams. 

This Flora of Guatemala, while prepared as a descriptive account 
of the plants of this country alone, will be found useful over a much 
wider area. In it are included most plants known at present from 
El Salvador, Honduras, and Nicaragua. It will be found to contain 
also the larger part of the species of the southern states of Mexico, 
Chiapas and Tabasco especially, and a large proportion of the plants 
of even more remote states. Additions and corrections to the Flora 
are being assembled as they come to light. Suggestions and com- 
ments are welcomed from outsiders, and will be duly considered for 
the future elaboration of parts of the Flora. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 7 

Due to their large size in comparison with the size of other families 
treated, the Orchidaceae have been published as a unit consisting of 
two separate numbers, while the Gramineae have been published as 
part II of the Flora. The present part I, beginning with Cycadaceae 
and ending with Bromeliaceae, therefore, includes all families of 
Monocotyledons not previously published, but excludes those mono- 
cotyledonous families already published in part III, part II (Grami- 
neae), and the two separate parts devoted to Orchidaceae. 



ACKNOWLEDGMENTS 

Almost every botanical publication is the result not of the labor 
performed individually by its author but of the cooperation over a 
long period of years of a large number of persons, ranging from 
learned scientists, some of whom date back to the time of Linne" or 
even earlier, down to the most humble and illiterate peasants. It 
would be preposterous for any one person at the present time to 
claim full personal credit for such a piece of work. It is, indeed, 
questionable whether some of the humblest of the cooperatives, 
whose names never appear in print, do not often deserve prime 
credit for their aid. 

At any rate, a very large number of people have cooperated, over 
a hundred years and more, to make possible the present account of 
the flora of Guatemala. So far as the botanists are concerned, they 
all must have enjoyed their work, else they need not have been doing 
it. Very few botanists ever are forced to travel to foreign lands; 
rather, they fight for the privilege. 

It is hard to believe that there is any normal person in North 
America or Europe who would not find something to interest him in 
Guatemala. Every one would find some phase of nature or man in 
Guatemala that would be new to him and should therefore hold his 
interest for at least a fleeting instant. Beauty, perhaps fortunately, 
can not be measured by any scientific standard, but if it could, Gua- 
temala would have few superiors in natural beauty. Nowhere are 
there more beautiful and majestic mountains, lovelier lakes and for- 
ests, more beautiful wild and cultivated flowers. Nowhere is there a 
climate that is more agreeable and more invigorating. Nowhere 
will one find more interesting, more highly varied, and more pictur- 
esque people than in Guatemala. 

The people of Guatemala have contributed much to the accumu- 
lation of data on which our Flora is based. From the highest to the 



8 FIELDIANA: BOTANY, VOLUME 24 

most humble they have been exceedingly helpful when informed of 
the purpose in our work of collecting and studying the flora of their 
country. This is no surprise to one familiar with the people of all 
Central America. It would be a surprise and at once arouse painful 
speculation, if the people were otherwise than helpful or agreeable. 
From the people of pure and mixed Spanish blood the greatest cour- 
tesy and consideration are to be expected as a matter of course; it is 
a racial trait. In Guatemala the Indians and the poor among the 
ladinos yield nothing in breeding to their social superiors. We have 
found them invariably dignified, courteous, kindly, and hospitable. 
Some of their standards of kindliness as exhibited among themselves 
may seem different from ours, but it must be understood that the 
poor of Central America often fight to exist under the most stern and 
harsh conditions. What at first glance may seem harsh to us need 
not be so for people actually concerned in the matter. A northerner 
may wince and shudder when he sees the heavy work performed by 
small children, yet let him observe the care with which a father 
watches over them, and he will realize that there is no intentional 
cruelty. Certainly no people are more tender and just to their chil- 
dren than the Indians of Guatemala. 

We prefer the Central Americans to help us when we go collect- 
ing. We have had many dozens of them to help in collecting, and 
so far as their ability and understanding of the work went, they were 
beyond criticism. 

It is quite out of the question to mention here all the persons to 
whom we are indebted for assistance in our field work in Guatemala, 
their help often of a most practical and frequently very substantial 
nature. We are sure that they do not expect thanks beyond those 
offered at the time their services were performed. There are a few, 
however, whom it would be unfair to omit, and mention is made 
below of at least some of them, the order in which they are named 
being purely casual. 

We are particularly indebted to a great number of officials of the 
Guatemalan government for their attentions and assistance in the 
course of our work. We received most substantial help in our work 
from thejefes politicos, director es de policia, and intendentes in all parts 
of the republic. Special thanks are due also to the engineers and their 
assistants of the national roads system, who always are helpful to 
travelers who encounter trouble along the roads. In some instances 
they supplied transportation for us and our baggage. 

We are especially grateful to Don Mariano Pacheco Herrarte, 
then Director General de Agricultura, who always has been a kind 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 9 

host to visiting scientists, and was particularly gracious to us with 
advice and practical assistance. Don Mariano's orchid garden is one 
of the favorite spots of interest to tourists who visit Guatemala, and 
it is of double interest to botanists. At the Feria Internacional, held 
each November in Guatemala City, his exhibits of orchids and agri- 
cultural products have been one of the most entertaining and educa- 
tional displays. One would never suppose ordinarily that a small 
country could produce so vast a variety of agricultural products. 
The agricultural exhibits are only part of this great fair, which is 
well worth a visit even if one has to travel a long distance to reach it. 

We are deeply indebted also to Professor Ulises Rojas, well- 
known botanist of Guatemala and formerly Director of the Jardin 
Botanico, who has trained most of the younger generation of scien- 
tists. He is celebrated also for his Elementos de botanica general, 
whose three volumes form a textbook of botany and contain a great 
deal of useful information regarding plant life of Central America. 
Professor Rojas has traveled widely in Guatemala and outside the 
country, in regions little or not at all known to ordinary visitors. 
During his travels he has gathered a wealth of information regarding 
the botany of the region, some of which he has kindly shared with us, 
thus greatly enriching the present Flora. The junior author was his 
guest for some weeks at his Finca Los Pirineos, from which he ex- 
plored the mountains of Quezaltenango and the mountain forest of 
the barranco of the Rio Samala. 

We are under obligations, too, to Don Jose" Ignacio Aguilar G., 
who during our visits was Director of the Finca Nacional La Aurora, 
on the edge of the capital. He later was instrumental in building 
there the new museum of natural history. Don Ignacio has been 
very helpful to us in many ways. He assembled a large series of plant 
specimens, more than 2,000 of them in fact, illustrating the flora of 
such departments as Guatemala, Escuintla, and Quiche", and this 
collection has been invaluable in preparation of the present Flora. 
He also furnished lists of the vernacular names applied to many of 
the plants. For many of these names he is cited as authority on the 
following pages. 

Others to whom the junior author is specially indebted for favors 
that greatly facilitated his Guatemalan work are the following: Gen- 
eral Aurelio F. Recinos, of Huehuetenango; Don Antonio Rodriguez, 
manager of the chrome mines near Jalapa; the late Don Alejandro 
Cordoba, proprietor of the newspaper El Impartial, who provided 
accommodations for collecting on his properties in Sierra de las 
Minas; Don Herman and Dona Mercedes Paz R., of Finca Cubilgiiitz 



10 FIELDIANA: BOTANY, VOLUME 24 

in Alta Verapaz; and Don Benvenuto Lopez, administrador of Finca 
Yalpemech, in the same department. 

The senior author wishes to express here his deep appreciation of 
the many courtesies extended to him by Don Guillermo Bonifaz and 
his charming family of Quezaltenango. The many weeks spent in 
that city are remembered with particular pleasure because of their 
cordial reception and kindly hospitality. 

We received much direct assistance, as we always have done in 
other parts of Central America, from the officials of the United Fruit 
Company, and in western Guatemala from the Compafiia Agricola 
de Guatemala. They provided ample facilities, otherwise nearly or 
quite impossible to obtain, for collecting in the lowlands where the 
banana plantations are located. We wish to mention especially in 
this connection Dr. N. C. Mcphail, of Quirigud Hospital, who is 
widely esteemed and respected in Central America; Mr. George Aus- 
tin, of Puerto Barrios, whose courtesy and patience with visiting 
scientists from the North seem never to be exhausted; and Dr. Wil- 
son Popenoe, who as always has been uncommonly helpful with his 
advice, the result of many years spent in Central America, and with 
his very substantial and direct aid in procuring quarters from which 
field work was done. Several officials of the Ferrocarril Internacional 
de Centro-Ame>ica also have extended favors that helped in the suc- 
cess of our work. We are particularly indebted to Mr. and Mrs. 
Bevan B. Lewis, formerly of Guatemala City. 

Others whom we wish to mention here, in recognition of their 
courtesies and their help in our field work, are the following: Mrs. 
Gordon Smith, of Finca Moca, Suchitepe"quez; Mr. Owen Smith, of 
Finca Panama in Solola; Mrs. R. W. Hempstead, of Coban; Mr. 
William E. Diesseldorff, of Coban; the late Mr. Enrique Zoller, for- 
merly manager of Finca El Porvenir, San Marcos; the late Dr. and 
Mrs. Edward Haymaker, of El Rancho; Mr. Frederic Rosengarten 
and Mr. Hoehne, of Finca El Naranjo; Mr. and Mrs. L. Lind Peter- 
sen, of Finca El Zapote, Escuintla; and Mr. Federico Kong 0., of 
Guatemala City. 

From botanists of the United States we have received a great 
deal of valued help in determination of certain Guatemalan groups. 
These persons are mentioned under the families or genera of plants 
in which they are interested. 

For data regarding the properties of Guatemalan woods we have 
relied on Timbers of the New World, by Samuel J. Record and Rob- 
ert W. Hess. For many years the late Professor Record and the 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 11 

senior author of this Flora were closely associated in studies of Cen- 
tral American forests, and often were able to be mutually helpful in 
the work. Professor Record's keen interest in all that pertained to 
the forests of Central America will be greatly missed. The volume 
of which mention is made here will be found invaluable to all who are 
in any manner interested in tropical American woods, since it is a 
veritable encyclopedia of information regarding all their phases. 

The senior author is exceptionally indebted to one person who 
during both his visits to Guatemala was a dependable counsellor and 
one of the most delightful traveling companions that ever could be 
found: Dr. John R. Johnston, then Director of the Escuela Nacional 
de Agricultura at La Alameda, near Chimaltenango. Several weeks 
were spent with great profit at La Alameda while enjoying Dr. John- 
ston's hospitality, and our many joint collecting trips will be long 
remembered. His enthusiasm in the field, his unfailing patience and 
his good humor in the face of petty difficulties, his keen interest in 
everything new, made field trips in his company a delight. His long 
years of experience in the Latin American countries have given him 
unusual appreciation of the countries and people of Central America, 
and few indeed are the North Americans who consider the Central 
American people with such sympathy and high regard. 

For the photographs from which the illustrations in this part of 
the Flora are made, we are indebted to the following persons: Dr. 
Albert E. Vatter, Jr., who accompanied the junior author on his sec- 
ond Guatemalan expedition, and made several hundred photographs 
in black and white and also in kodachrome; to Dr. John R. Johnston, 
who made many photographs while Director of the Escuela Nacional 
de Agricultura; and to Dr. C. L. Wilson, of Dartmouth College, who 
has made many handsome photographs in the lower mountains of 
Alta Verapaz. 

CYCADACEAE. Cycad Family 

Reference: J. Schuster, Cycadaceae, Pflanzenreich IV. 1: 1-168. 
1932. 

Palm-like plants, the caudex sometimes bulb-like or tuber-like, often columnar 
and elongate, simple or branched, bearing at its apex few or numerous leaves; leaves 
spirally arranged, often forming a dense crown at the apex of the trunk, coriaceous, 
pinnate or bipinnate, the leaflets opposite or alternate, usually linear or lanceolate, 
often oblique, glabrous in age but often tomentose when young, entire or dentate, 
parallel-nerved, the petiole usually aculeate; flowers dioecious, naked, the inflores- 
cences borne at the apex of the caudex among the leaves, strobiliform, terminal or 
axillary, the staminate strobiles narrower than the pistillate; microsporophylls 
thick, spreading at about a right angle, short-stipitate, narrowly or broadly cune- 



12 FIELDIANA: BOTANY, VOLUME 24 

ate, bearing on the lower surface very numerous microsporangia, the terminal 
portion sterile, acuminate, attenuate, or peltate; microsporangia sessile or very 
shortly stipitate, 2-6 aggregate in a sorus, oblong or subglobose, dehiscent at the 
apex by a median line; pollen smooth, ellipsoid or subglobose; pistillate strobiles 
pedunculate, cylindric or ovoid; macrosporophylls various in form, pedicellate, the 
terminal portion usually peltate-dilated, the macrosporangia inserted at the base of 
the peltate portion; macrosporangia orthropous, sessile or subsessile, covered by 
a single tough integument, the micropyle long and narrow; seeds oval or broadly 
ellipsoid or globose, the putamen smooth, striate, or costate, terete or obtusely 
angulate, usually with osseous foramina at the base; endosperm fleshy; cotyledons 
2, rarely 3-6, fleshy, oblong. 

Nine genera are recognized, of rather wide but often localized 
distribution in the tropics of both hemispheres. Only four of the 
genera are American, and only three genera are represented in Cen- 
tral America. 

Leaflets 1-nerved, not articulate with the rachis Cycas 

Leaflets several-nerved, articulate with the rachis. 

Scales of both pistillate and staminate cones with 2 horn-like appendages at the 
center Ceratozamia 

Scales of the pistillate and staminate cones peltate, not appendaged Zamia 

CERATOZAMIA Brongniart 

Trunk low, ovoid or cylindric, simple or dichotomously branched; petioles 
aculeolate, the rachis of the leaf circinnate in vernation, the undeveloped leaflets 
imbricate; leaflets elongate, narrowly or broadly lanceolate to obovate, attenuate- 
acute, narrowed to the base, opposite or alternate, articulate at the base, entire, 
the margins revolute, several-nerved; staminate strobile cylindric or narrowly 
elongate; microsporangia cuneiform, the sterile portion pyramidal-truncate or sub- 
applanate, with 2 horn-like appendages; pistillate cone ellipsoid; macrosporophylls 
ovuliferous on both sides, borne on a stout stipe, the apex transverse-hexagonal, 
bearing 2 horn-like appendages at the middle; macrosporangia ovoid, sessile. 

Two species are usually recognized, one Mexican and apparently 
little known. Chamberlain recognizes at least six species. 

Ceratozamia mexicana Brongn. Ann. Sci. Nat. III. 5: 8. pi. 1. 
1846. Costilla de Ie6n. Figure 1. 

Rocky wooded stream banks or rocky hillsides, 900-1300 meters; 
Huehuetenango (Sierra de los Cuchumatanes: San Andre's; Finca 
Soledad, 5 miles southeast of Barillas; Rio Amelco). Oaxaca and 
Veracruz, and perhaps elsewhere in southern Mexico. 

Plants acaulescent or with a short, ovoid or globose trunk, in age becoming 
longer and cylindric, covered by the persistent cataphylls and leaf bases, more or 
less fuscous-tomentose, simple or sometimes dichotomous; leaves erect at first, 
spreading in age and recurved, 1-3 meters long, the young ones often pruinose- 




S.H. Or ov . 



FIG. 1. Ceratozamia mexicana. a, Staminate strobile (X }). b. Cone scale 
(X M). c, Habit, pistillate plant ( X ^). 



13 



14 FIELDIANA: BOTANY, VOLUME 24 

glaucous and pilose, the larger ones with 7-75 pairs of leaflets; petioles terete, 
armed with short sharp spines for their whole length, the rachis armed with scat- 
tered small spines; leaflets alternate or opposite, straight or slightly falcate, nar- 
rowly or broadly lanceolate or subdimidiate-obovate from a cuneate base, gradually 
or abruptly attenuate-acute, the margins thick and subrevolute, paler beneath, 
coriaceous, 14-35 cm. long, 2-7 cm. wide, 17-40-nerved; staminate strobile cylin- 
dric, acute or subobtuse, 50 cm. long or shorter, 1.5-5 cm. thick, pedunculate; 
microsporophylls cuneiform, 1-1.5 cm. long, the fertile portion 1 cm. long; pistil- 
late strobile ellipsoid, conic-acute, 8-15 cm. long, 1.5-6.5 cm. broad, borne on a 
peduncle 5 cm. long; cone scales hexagonal at the apex, 1.5-2.5 cm. broad, fuscous- 
tomentose; putamen ligneous, subellipsoid, substriate, brownish, about 2 cm. long 
and 1 cm. broad. 

A showy and handsome plant, better adapted to cultivation than 
the local species of Zamia. Plants brought from Finca Soledad are in 
cultivation in the garden of Don Marcos Recinos at Huehuetenango. 



CYGAS L. 

Plants usually tree-like, the trunk cylindric, erect; leaves pinnatisect, the leaf- 
lets numerous, the lowest ones often spine-like, linear or linear-lanceolate, the 
margins incurved or thickened; staminate strobile oblong-ovoid or oblong, the 
microsporophylls cuneiform, usually acuminate; pistillate strobile terminal in the 
center of the trunk; macrosporophylls imbricate, borne on a linear stipe, the blades 
various in form, bearing on either side near the narrowed base 2 or more alternate 
or opposite macrosporangia, these ovoid-globose or obovoid; putamen more or less 
biangulate or rarely triangulate. 

Five species are usually recognized, in the Old World tropics. 
Chamberlain recognizes at least twenty species. Sago starch ob- 
tained from the trunks has been and still is an important food in 
regions where the plants are native. The seed coats have a good 
percentage of oil and are rarely used as food. The starch-rich ker- 
nels, however, are commonly used as food. 

Gycas revoluta Thunb. Fl. Japon. 229. 1784. 

Planted commonly for ornament in parks and gardens of Guate- 
mala, from the Pacific coast up to the central highlands (to at least 
1500 meters), also about Coban, and doubtless in other parts of the 
country. Native of the East Indies and southeastern Asia. 

Adult trunks sometimes attaining a height of 8 meters but usually lower, often 
very thick, subannulate, the leaf scars rhombic-quadrangular; leaves often very 
numerous, borne at the top of the trunk, about 75 cm. long, long-persistent, tomen- 
tose at first but soon glabrate, the petioles spiny; leaflets about 125 pairs, crowded, 
the largest ones 9-18 cm. long and 5-6 mm. wide; staminate strobile cylindric or 
ovoid-oblong, 8-40 cm. long, 1.5-4 cm. thick; seeds yellow-orange or yellow, cov- 
ered with a dense tomentum, 1.5-3.5 cm. long. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 15 

There are many handsome individuals of this species in the parks 
of Guatemala, as in the Parque Central of Guatemala, in Antigua, 
Coban, and Retalhuleu and Mazatenango. Cones are often pro- 
duced. Rare in cultivation in Guatemala is Cycas circinalis L., of 
the East Indies, in which the leaflets are about 12 mm. wide, and 
have flat rather than incurved margins. 



Dioon edule Lindl., a Mexican plant with pale foliage and very 
numerous rigid, not articulate leaflets, is in cultivation in Guatemala, 
but is rare. The genus Dioon, consisting of three or four Mexican 
species of tree-like plants, extends southward in Mexico to the State 
of Chiapas. As a species, D. Mejiae Standl. & Williams, has recently 
been described from Honduras; it is probable that the genus occurs 
in Guatemala. The rather ambiguous and little-known D. Dohenyi 
E. A. Howard has been ascribed through some mistake to Guate- 
mala, but its author and collector states (Touring Topics, March, 
1933, p. 11) that it was actually collected in the mountains of Chia- 
pas, about 100 miles north of the Guatemalan border and 30 miles 
inland from the coast. 

ZAMIA L. 

Plants usually low and with a hypogaean trunk, or sometimes developing above 
ground an elongate caudex, often tuber-like, simple or sparsely branched; leaves 
pinnate, with few or numerous leaflets, the petiole aculeate or unarmed, the leaflets 
articulate at the base, entire or dentate near the apex, mostly linear or lanceolate; 
staminate strobiles cylindric or elongate-cylindric; microsporophylls pedicellate 
and peltate or cuneiform-peltate, the microsporangia arranged beneath in 2 separate 
or confluent areas or in a marginal series on each side; scales truncate-pyramidal 
to hexagonal; pistillate strobile cylindric or ovoid-elongate; macrosporophylls 
pedicellate-peltate, biovulate beneath, the scales hexagonal or subquadrangular; 
macrosporangia ovate or subtrigonous-ovate, ferruginous to orange, yellow, or 
whitish. 

A group of about 25 species, all American, chiefly in North Amer- 
ica and West Indies. The Central American species still are imper- 
fectly known because of the scant material available for study, but 
about eight species have been reported from all Central America. 

Leaflets entire, rarely with a few teeth near the apex. 

Leaflets obovate to broadly oblanceolate, mostly 6-10 cm. wide . Z. Tuerckheimii 
Leaflets lance-linear, mostly 2-3 cm. wide Z. monticola 

Leaflets conspicuously spinulose-serrate, usually from about the middle to the 
apex, sometimes for almost their whole length. 

Leaflets relatively small, mostly 1.5-2.5 cm. wide Z. Loddigesii 

Leaflets larger, mostly 5-7 cm. wide Z. muricata 



16 FIELDIANA: BOTANY, VOLUME 24 

Zamia Loddigesii Miq. Tijdschr. Nat. Geschied. 10: 73. 1843. 
Z, latifolia Lodd. ex A. DC. in DC. Prodr. 16, pt. 2: 541. 1868. Z. 
cycadifolia Dyer in Hemsl. Biol. Centr. Amer. Bot. 3: 195. 1883. 
Z. Herrerae Cald. & Standl. Journ. Wash. Acad. Sci. 14: 93. /. 1. 1924. 
Z. spartea A. DC. op. cit. 539. 1868. Z. Loddigesii var. longifolia 
Schuster, Pflanzenreich IV. 1 : 147. 1932. Z. Loddigesii var. latifolia 
Schuster, loc. cit. Z. Loddigesii var. cycadifolia Schuster, op. cit. 148. 
Z. Loddigesii var. spartea Schuster, op. cit. 148. Camotillo; Cocalito; 
Teosinte (north coast, fide Rojas); Chacuhua (Pete"n, fide Lundell). 
Figure 2. 

Open slopes or fields, sometimes in wet or moist thickets or forest, 
occasionally in lowland pine forest, lowlands of both coasts at sea 
level or but little higher; Pete"n; Izabal; Santa Rosa; Retalhuleu. 
Southern Mexico and British Honduras; Honduras; El Salvador. 

Caudex usually hypogaean, subcylindric or often tuberous and fleshy, some- 
times 20 cm. high; petioles densely or sparsely aculeate, the spines short, straight, 
rigid, mostly 1-2 mm. long, puberulent or glabrate; leaflets variable in size and 
shape, linear-lanceolate to obovate-oblong, usually 20 pairs or fewer, long-attenu- 
ate to subobtuse at the apex, mostly 10-25 cm. long and 12-25 mm. wide, some- 
times somewhat larger, opposite or alternate, spinulose-serrate, usually conspicu- 
ously so, above the middle or for almost their whole length; staminate strobiles 
3.5-6.5 cm. long, about 2 cm. thick, conic or obtuse at the apex, the scales fulvous- 
tomentose, 4-6-sided; pistillate strobile thick-cylindric, cuspidate, about 5.5 cm. 
long, the peduncle equaling or often exceeding the strobile, its scales quadrangular- 
rhombic or 6-angulate, 7-8 mm. wide; seeds ligneous, ovoid, about 13 mm. long 
and 7 mm. wide. 

According to Schuster, the plant reported by Hemsley (Biol. 
Centr. Amer. Bot. 3: 194. 1883) as Z. Ottonis Miq. is perhaps refer- 
able here. The plant is a variable one, segregated by some authors 
into several species, and Schuster recognizes several varieties, of 
which vars. longifolia and spartea are reported from Guatemala, but 
the varieties are based upon what appear to be characters of scant 
importance. The plant is sometimes seen in Guatemalan gardens 
and is said to grow readily from seeds. It has been reported from 
Guatemala under the name Z. furfuracea L., which pertains to a 
Mexican species. The camotillo is one of the most celebrated plants 
of Central America, where it is widely known because of its poisonous 
properties. The large starchy roots are reported to be used in Pete"n 
for poisoning rats, and it is believed that they have been used many 
times in Central America for criminal poisoning of human beings. 
There is a popular belief that if the roots have been out of the ground 
one day and are used for poisoning, they kill in one day; if out of the 
ground ten days, they kill in ten days, and so on. It is unnecessary 




FIG. 2. Zamia Loddigesii. Habit (X 
17 



18 FIELDIANA: BOTANY, VOLUME 24 

to state that there can be little basis for this belief, but there is no 
doubt that they are poisonous. The starchy roots of various species 
of the genus were much used by some of the aborigines, especially 
the Caribs, as food, but they well knew that the poisonous properties 
must first be removed by thorough cooking. Some of the Zamia 
species supplied the article known as Coontie, an important food 
staple of the Florida Indians. Wisdom reports that in the Jocotan 
region of Chiquimula the dry roots of what are presumed to be iden- 
tified as Zamia are sometimes chewed and the saliva is swallowed to 
relieve cough and also to improve the singing qualities of the voice. 
He states further that the roots mixed with lime are used by the 
Indians of Jocotan for coloring tobacco pipes and other articles red. 
The name "cocalito" was said to be given this plant because of the 
resemblance to a small coconut palm, but the resemblance is not 
particularly striking. 

Zamia monticola Chamberlain, Bot. Gaz. 81: 219. /. 1-13. 1926. 

Alta Verapaz, cliffs above Rio Chiacte", 540-600 meters, on lime- 
stone, Finca Volcan; also at Finca Arenal; between Chajmayic and 
Sebol, at 300-500 meters. Type from Volcan de Naolinco, near 
Jalapa, Mexico. 

Trunk as much as 20 cm. high and 15 cm. in diameter; petioles sparsely acule- 
ate; leaves as much as 2 meters long, the leaflets about 30-34 pairs, opposite or 
alternate, linear-lanceolate, falcate, 24-35 cm. long or more, 3-4 cm. wide, gradu- 
ally attenuate to the tip, shortly narrowed at the base, lustrous when dried, 
sometimes slightly serrulate near the apex but usually entire, the nerves about 30; 
staminate strobiles oblong-ovoid, 12-16 cm. long, on a peduncle 10-17 cm. long; 
microsporophylls with a hexagonal apex; microsporangia 10-16, in 2 widely sepa- 
rated series. 

Zamia muricata Willd. Sp. PL 4: 847. 1805. Z. variegata Warsc. 
in Otto & Dietr. Allgem. Gartenz. 13: 253. 1845. Z. picta Dyer in 
Hemsl. Biol. Centr. Amer. Bot. 3: 194. 1883. Z. muricata var. picta 
Miq. Wiss. Nat. Tijdschr. 1: 198. 1846-47. Camotillo. 

Wet forest or dry pine ridges, Izabal, at 600 meters or less. Re- 
ported also from Mexico (Oaxaca), Colombia, and Venezuela. 

Trunk sometimes 2.5 meters tall, often wanting, at least in young plants, 
simple or branched; leaves large, the petiole terete, aculeate, glabrous except at 
the base, there tomentose; leaves 10-12 pairs or more, alternate or subopposite, 
lanceolate or oblong-lanceolate, sometimes oblong-oblanceolate, up to 35 cm. long, 
sometimes 8 cm. wide, acuminate, contracted at the base, spinulose-denticulate 
from the apex sometimes almost or quite to the base, lustrous above when dried, 
paler beneath; staminate strobiles cylindric, conic-acuminate or obtuse, 3-6 cm. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 19 

long, 1 cm. thick, on peduncles 4-11 cm. long, the scales hexagonal, 6 mm. broad 
or less; pistillate strobiles cylindric, 5-11 cm. long, 3-3.5 cm. thick, on a peduncle 
4-6 cm. long; seeds ovoid, trigonous-compressed, red at maturity, 3 cm. long, 
2 cm. thick. 

Var. picta was based upon plants grown in Europe and supposed 
to be of Guatemalan origin. There is some question as to the deter- 
mination of the recent collections referred to this species. They may, 
in fact, represent two distinct species. They are closely related to 
Z. Skinneri Warsc., so far as leaves are concerned, except that the 
leaf nerves are not sharply elevated as in that plant of Panama and 
Costa Rica. It seems very improbable that a species of this genus 
actually ranges from Guatemala to Venezuela, especially since it has 
not been collected in the intervening areas. Further and more com- 
plete collections are necessary to determine the relationship of all the 
Guatemalan members of the genus Zamia. Most of the recent col- 
lections are sterile, although it must be stated that the leaves and 
not the cones seem to afford the most dependable characters for sep- 
arating the species, the cones apparently varying but little in closely 
related species. 

Zamia Tuerckheimii Bonn. Smith, Bot. Gaz. 35: 8. pi. 1. 1903. 
Figure 3. 

Wet limestone thickets or forest, Alta Verapaz, 300-1100 meters; 
type from Cubilgiiitz, Tuerckheim 7786; collected also at Finca Vol- 
can, where said to be common on slopes of the higher hills; at Finca 
Seamay, near Senahu; and on rocks between Campur and Socoyo, 
on the Pete'n Highway. Reported from British Honduras; Atlantic 
coast of Honduras (Lancetilla Valley near Tela). 

Trunk almost none or often elongate and sometimes 3 meters long, often 
pendent from rocks and S-shaped, 20 cm. in diameter or less, simple; petioles slen- 
der, sparsely aculeate or sometimes unarmed; leaflets mostly 5-9 pairs, subopposite, 
very lustrous when dried, somewhat paler beneath, ovate-lanceolate to oblong- 
elliptic or oblong-obovate, mostly 12-25 cm. long and 4-8 cm. wide, abruptly 
acuminate, entire, contracted at the base, the nerves 40 or more; staminate strobile 
narrowly cylindric, about 14 cm. long and 2.5 cm. wide, apiculate, the scales trun- 
cate-pyramidal, hexagonal; pistillate strobile borne on a peduncle 2 cm. long, about 
18 cm. long and 6 cm. broad; seeds red, obovoid, 2 cm. long, 13 mm. broad. 

Excellent material of the species was collected around Finca Vol- 
can by Dr. C. L. Wilson, who also made handsome photographs of 
the plant. It is plentiful in some places along the newer part of the 
Pete'n Highway, growing upon the tops and sides of great jagged 
blocks of the characteristic limestone of the region. The S-shape of 
the more elongate trunks is very characteristic of the plant. 



20 



FIELDIANA: BOTANY, VOLUME 24 





FIG. 3. Zamia Tuerckheimti. a, Pistillate strobile (X }>) b, c, Macrosporo- 
phyll (X 1 A). d, Macrosporangium (X M). e, Habit of pistillate plant (X Vie). 
/, Portion of leaf (X Y). g, Habit of staminate plant (X Vw). h, Staminate 
strobile (X Yi}. i, Microsporophyll, viewed from above (X Ijiz). j, k, Micro- 
sporophyll (X 



PODOCARPACEAE 

Reference: R. Pilger, Taxaceae, Pflanzenreich IV. 5: 94. 1903. 

Usually tall trees; leaves usually spirally arranged, rarely opposite, persistent, 
thick and coriaceous, often spreading from the branch in a single plane, linear or 
lanceolate, rarely ovate, sometimes scale-like; flowers usually dioecious; staminate 
flowers composed of stamens only, sometimes surrounded at the base in bud by 
involucre-like rigid scales, terminal or solitary and axillary or in few-flowered, 
mostly axillary inflorescences, these usually ament-like, the stamens numerous, 
spirally imbricate; anthers 2-celled, usually ovoid and dehiscent by slits; pistillate 
flowers usually solitary in the leaf axils, the flowers naked, often subtended by 
sterile scales; carpels 1-many, each consisting of a single naked ovule, usually con- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 21 

nate with an epimatium (an excrescence from the carpel); fruit larger than the 
flowering carpel but retaining its form, the seed covered by a coat consisting of the 
integument and the enlarged epimatium. 

About 5 genera, in tropical and temperate regions of both hemi- 
spheres. A single genus occurs in North America. 

PODOCARPUS L'Hentier 

Reference: John T. Buchholz and Netta E. Gray, A taxonomic 
revision of Podocarpus, Journ. Arnold Arb. 19: 123-151. 1948. 

Usually tall trees; leaves linear to lanceolate or ovate, usually acute or mucro- 
nate, spirally arranged or rarely opposite, usually spreading in a single plane from 
the branch; flowers dioecious or rarely monoecious, the staminate usually solitary 
or few and sessile or pedunculate in the leaf axils, surrounded at the base by sterile 
scales; anthers usually imbricate, 2-celled; pistillate flowers usually solitary and 
pedunculate in the leaf axils, the receptacle fleshy, with 1-2 carpels, these 1-ovu- 
late; seeds usually large, more or less apiculate, enclosed in a double testa, the outer 
fleshy, the inner usually ligneous. 

About 60 species, in tropical or temperate regions of both hemi- 
spheres. One other species and several additional varieties are 
known from southern Central America (Costa Rica and Panama). 
Podocarpus Blumei Endl., native of southern Japan, is in cultivation 
in Finca La Aurora, Guatemala. It has ovate to lance-ovate leaves 
as much as 3-4 cm. wide. Another species, P. Harmsianus Pilger, 
native to South America, and similar to P. Standleyi Buchholz & 
Gray of the Costa Rican mountains, is planted in the garden of Don 
Mariano Pacheco in Guatemala. 

Terminal bud scales long-acuminate, 3-4 or more times as long as wide .P. Matudai 
Terminal bud scales ovate, acute, obtuse or apiculate, but not attenuate. 

Vegetative bud scales apiculate P. guatemalensis 

Vegetative bud scales acute or obtuse P. oleifolius 

Podocarpus guatemalensis Standl. Proc. Biol. Soc. Wash. 37: 
49. 1924. 

Known only from the type, Puerto Barrios, Dept. Izabal, Stand- 
ley 25090. 

A tree up to 20 meters tall, the trunk up to 70 cm. in diameter; bark reddish 
brown, scaly; vegetative buds spherical or ovoid with broad ovate bud scales 
apiculate or obtuse, scarious at margins or not so; leaves lanceolate, those of lower 
branches or saplings up to 14 cm. long, 10-12 mm. wide, distinctly ridged at mid- 
vein, flat or depressed below midvein, cuneately narrowed near base, narrowed 
toward apex, those of mature reproductive branches usually up to 10 cm. long and 
less than 10 mm. wide; upper hypoderm interrupted; auxiliary sclereids lacing in 



22 FIELDIANA: BOTANY, VOLUME 24 

palisade or mesophyll; vascular sclereids lacking; pollen cones cylindrical, axillary 
to leaves or scales of previous year's growth; seed cones axillary; seed on short 
peduncles up to 4-5 mm. long, receptacles up to 7 mm. long; seeds ellipsoidal, up 
to 8 mm. long with small conical crests. 

The type was taken from a low shrub growing in the densely 
wooded swamps about Puerto Barrios. The shrub was probably a 
casual introduction there, perhaps brought down from the hills by 
some stream, for further exploration in the neighborhood has failed 
to reveal other plants of the species, which grows normally in the hills. 

Two varieties have been recognized, Podocarpus guatemalensis 
var. pinetorum (Bart.) Buchholz & Gray, known only from British 
Honduras, and P. guatemalensis var. Allenii (Standley) Buchholz & 
Gray, known from Costa Rica and Panama. Podocarpus guatemalen- 
sis var. pinetorum, the type of which comes from Mt. Pine Ridge, 
El Cayo District, Bartlett 13109, differs from typical P. guatemalensis 
in having much larger terminal buds with broader, thicker scales and 
shorter, narrower, and thicker leaves. 

Podocarpus Matudai Lundell, Phytologia 1: 212. 1937. 

Forests of southern Mexico at 1250-2370 meters, the type from 
Mt. Pasitar, Chiapas, Matuda 698. 

A large tree, the trunk up to 1.5 meters or more in diameter; terminal buds 
large with narrowly lanceolate, somewhat spreading scales; leaves coriaceous, 
lanceolate, 4-9 cm. long, 10-15 mm. wide, up to 16 cm. long and 19 mm. wide on 
vigorous shoots, narrowly acute at apex, narrowed to a petiole below, with a some- 
what rounded ridge above midvein, becoming less prominent toward apex and less 
prominent below; auxiliary sclereids scattered in palisade and mesophyll, vascular 
sclereids both above and below bundle; pollen cones sessile, cylindrical, arising 
from axillary buds of previous season's growth, becoming 3-3.5 cm. long, 4 mm. 
wide, surrounded by broad, carinate apiculate scales, shedding pollen in January 
or later; microsporophylls with upturned obtuse apices with scarious, denticulate 
margins; seed cones on short peduncles 4-6 mm. long, receptacle of a pair of fused 
thickened scales 4-6 mm. long, with short obtuse apices; seed 8-10 mm. long, 
7-8 mm. wide, with a blunt crest scarcely evident. 

Podocarpus Matudai var. macrocarpus Buchholz & Gray, 
Journ. Arnold Arb. 19: 132. 1948. Ciprecillo; Tabla (Chiapas); 
Curus-te (Huehuetenango) . 

Usually in moist or wet, mountain forest, mostly at 1200-2600 
meters, but sometimes occurring much lower in Guatemala; El Pro- 
greso (Sierra de las Minas, Montana Canahui); Chiquimula (Cerro 
Brujo) ; Guatemala; Huehuetenango (at various places in Sierra de 
los Cuchumatanes; between Ixcan and Finca San Rafael at only 
200-800 meters). Chiapas (type from Mt. Ovanda). 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 23 

Like the species, but with longer pistillate peduncles, 9-18 mm. long, and 
larger seeds, more than 12 mm. long; pollen cones 4-6 cm. long, maturing earlier 
than the species. 

Podocarpus oleifolius D. Don in Lambert, Pinus 2: 20. 1824, 
ed. 2. 2: 121. 1828. 

Cloud forests at 2000-3200 meters in Guatemala; Zacapa (Sierra 
de las Minas, Cerro de Monos and Rio Repollal, Volcan Gemelos) ; 
Baja Verapaz (Sierra de las Minas east of Chilasco) . Southern Mex- 
ico (Chiapas and Oaxaca); El Salvador; Honduras; Costa Rica; 
western South America. Figure 4. 

A tree up to 20 meters tall; bark yellowish brown; vegetative buds globose, 
the bud scales rounded or broadly ovate, obtuse and scarious margined; leaves 
coriaceous, narrowly lanceolate, 2.5-8 cm. long, 5-12 mm. wide, up to 15 cm. long 
and 20 mm. wide on sprouts and vigorous shoots, with a conspicuous narrow groove 
above the midvein, broad but not very prominent beneath; upper hypoderm con- 
tinuous; vascular sclereids usually present; auxiliary sclereids usually present; 
pollen cones solitary, axillary, 3 cm. long, 3 mm. wide, arising from sessile buds of 
previous year's growth; microsporophylls tipped with upturned rounded apiculus; 
pistillate cones solitary, axillary, the peduncles 5-10 mm. long, receptacle of 2 fused 
fleshy scales free at tip, 6-9 mm. long; seed globose-ovoid with a suppressed crest, 
7-8 mm. long. 

Three varieties are recognized, two (var. macrostachyus and 
var. trujillensis) from northern South America and one (var. costa- 
ricensis) from Costa Rica and Panama. 

ARAUCARIACEAE 

Reference: R. Pilger, Araucariaceae, Pflanzenfamilien ed. 2. 13: 
249-266. 1926. 

Mostly large trees; leaves small or large, needle-like or rather broad, persistent; 
flowers dioecious or rarely monoecious; staminate flowers large, terminal and fas- 
ciculate on short branchlets, or sometimes axillary; stamens numerous, spirally 
arranged, the filament dilated into a broad scale, the sporangia or anthers numer- 
ous, free, linear, arising on the lower side of the scale; pistillate flowers terminal on 
short branchlets, consisting of numerous scales without bracts; fruit a globose or 
ovoid cone, falling apart at maturity, the scales numerous, spirally arranged, 
broad, imbricate, thickened at the apex, sometimes winged; ovary 1; cotyledons 
usually 2, rarely 4. 

Only one other genus is known, Agathis, with about 20 species in 
southeastern Asia and the islands of the southern Pacific. All species 
of both genera are confined to the southern hemisphere. 

ARAUCARIA Jussieu 

Tall evergreen trees, the branches regularly verticillate, spreading; leaves alter- 
nate, decurrent at the base, spreading, sometimes 2-ranked, subulate to ovate- 





c H Gr'ove 



FIG. 4. Podocarpus oleifolius. a, Leafy branch with staminate flowers ( X 
b, Fruit (X%). 

24 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 25 

lanceolate, usually very stiff; flowers commonly dioecious, the staminate terminal 
and fasciculate; anthers with 8-15 sporangia; pistillate flowers terminal, ovoid or 
subglobose, composed of numerous scales without bracts; fruit a cone, ovoid or 
subglobose, falling apart when mature, the scales cuneate; seeds large, adherent 
to the scales; cotyledons 2-4. 

About 10 species, in South America, Australasia, and the Pacific 
Islands. Some of the species are important forest trees of South 
America, from Brazil southward, where they furnish timber used 
extensively for furniture and general construction. The large seeds 
of some species are edible. 

Leaves broad and flattened, 6-17 mm. wide, mostly 1.5-5 cm. long ... .A. Bidwllti 

Leaves all or mostly subulate or needle-like, the adult ones only 1-4 mm. wide and 

5-12 mm. long. 
Leaves of sterile branches 4-12 mm. long, curved upward, not spine-tipped, 

triangular in cross section A. excelsa 

Leaves of sterile branches 7-25 mm. long, almost straight, spine-tipped, much 
compressed A, Cunninghamii 

Araucaria Bidwillii Hook. Lond. Journ. Bot. 2: 503. 1843. 

Native of Australia; planted frequently in Guatemala for orna- 
ment. 

A large tree as much as 50 meters high, the trunk free of branches for half its 
length or more; leaves appearing 2-ranked on the branches, spreading, narrowly 
lanceolate to ovate-lanceolate or broader, spine-tipped, deep green and lustrous, 
very thick and rigid; branches usually in whorls of 10-15; staminate flowers 5-7.5 
cm. long; cones globose-ovoid, about 22 cm. long and 18 cm. broad, the scales 
2.5-5 cm. wide, with acute edges; seeds 5-6.5 cm. long, 2 cm. broad, with long 
compressed curved tips. 

There are numerous fine trees of this species planted in the moun- 
tains of Guatemala, chiefly at middle elevations but frequently as 
high as 2700 meters. Some of them have probably attained almost 
as great a height as those native in Australia. Large trees are to be 
seen in many of the parks and sometimes in the larger patios, particu- 
larly about Antigua, Totonicapan, and San Marcos. 

Araucaria Cunninghamii Sweet, Hort. Brit. 475. 1827. 

Native of Australia; planted for ornament about San Marcos, 
and doubtless in other parts of Guatemala. 

A tree sometimes 60 meters high in its native habitat, the branches in whorls 
of 4-7, the lower ones widely spreading, the upper branches ascending; leaves rigid, 
slender, almost acicular, sharp-pointed, straight or nearly so, strongly compressed 
laterally, conspicuously carinate on both sides, quadrangular, scarcely falcate; 
staminate flowers 5-7.5 cm. long; cones ovoid-globose, 7.5-9.5 cm. long, 7.5 cm. 



26 FIELDIANA: BOTANY, VOLUME 24 

broad; cone scales broadly cuneate, coriaceous, 1.2 cm. wide, terminating in a rigid 
recurved mucro. 

Araucaria excelsa R. Br. in Ait. Hort. Kew. ed. 2. 5: 412. 1813. 
Norfolk Island Pine. 

Native of Norfolk Island, between New Zealand and New Cale- 
donia; planted occasionally for ornament in Guatemala parks and 
gardens, chiefly at middle or rather high elevations. 

A tall tree, sometimes 60 meters high, at first pyramidal, in age with a more 
elongate crown, the branches in whorls of 4-7, horizontal or the lower ones some- 
what drooping; leaves small, subulate, upcurved, dark green, closely set on the 
branchlets; cones subglobose, brown, 10-15 cm. in diameter; cone scales 8-9 mm. 
long, with a stout incurved mucro. 

Seedlings of this tree make handsome pot plants, often seen in 
the United States and occasionally in Guatemala. One very large 
tree at the Hotel Manche"n in Antigua was said by its owner to be 
45 meters high, but the accuracy of the measurement is questionable. 
Great quantities of the terete older branchlets often are shed by the 
trees, sometimes completely covering the ground. 



CUPRESSACEAE. Cypress Family 

Reference: R. Pilger, Cupressaceae, Pflanzenfamilien 13: 361-403. 
1926. 

Trees or shrubs, the trees usually not very large, the shrubs sometimes pros- 
trate, often densely branched; leaves small, scale-like or needle-like but short, 
decussate or 3-ranked, most often closely appressed to the branchlets, both needle- 
like and scale-like leaves often found on the same plant, the leaves of seedling 
plants all needle-like; flowers monoecious or dioecious, very small, solitary and 
terminal on short twigs or solitary in the leaf axils, the sporophylls opposite 6r 
ternate; stamens with short filaments and broad anther-bearing scales; sporangia 
usually 3-6, ellipsoid, free; pistillate inflorescence of 3-8 scales, some or all of these 
bearing 1-2 ovules; scales of the pistillate inflorescence becoming woody or fleshy 
in fruit and more or less completely fused, or sometimes free at maturity; seeds 
small, free, sometimes winged; cotyledons usually 2, rarely 5-6. 

Sixteen genera are known, widely dispersed in both hemispheres, 
chiefly in temperate regions. Only two genera are native in Central 
America, and these only in Guatemala. Two others, bibocedrus and 
Chamaecyparis, are found in North America. 

Branchlets compressed and flat; fruit a woody dehiscent cone Thuja 

Branchlets terete or 4-sided. 

Fruit berry-like, fleshy, indehiscent, 1-4-seeded; needle-like leaves, when pres- 
ent, with whitish bands or marks on the upper surface Juniperus 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 27 

Fruit a dry dehiscent woody cone, many-seeded; needle-like leaves, when pres- 
ent, with whitish bands or marks on the lower surface Cupressus 

CUPRESSUS L. Cypress 

Evergreen trees or shrubs, the bark shredded, the branchlets terete or 4-angu- 
late; leaves aromatic, opposite, small and scale-like, minutely denticulate, ap- 
pressed, but on young plants or branches linear-subulate and spreading; flowers 
solitary, minute, terminating short branchlets, monoecious, the staminate and 
pistillate on separate branches; staminate flowers ovoid or oblong, yellow, consist- 
ing of 6-12 decussate stamens, each stamen with 2-6 pendulous globose anther 
cells; pistillate flower subglobose; cones globose or subglobose, maturing the second 
season, composed of 6-12 peltate scales, these becoming ligneous, usually with a 
mucro at the flattened apex of each scale, this bearing numerous, compressed and 
angulate, winged seeds, the lower scales usually smaller and sterile; cotyledons 3-4. 

Species about 12, in the warmer temperate and subtropical re- 
gions of both hemispheres, represented in Central America by a single 
species. In America the genus reaches the southern limit of its dis- 
tribution in Guatemala. The heartwood in this genus is yellowish, 
pale brown, or pinkish, sometimes streaked or variegated, the sap- 
wood white; of rather high luster, fragrant; light and soft to hard and 
heavy, of very fine and uniform texture and straight or irregular 
grain; easy to work and of high durability. The lasting properties 
of the wood are attested by the fact that wood of Old World cypress, 
Cupressus sempervirens L., was used for constructing the gates of 
Constantinople and the doors of St. Peter's, which remained sound 
until their removal after eleven centuries of use. 

Cupressus lusitanica Miller, Card. Diet. ed. 8. no. 3. 1768. 
C. Lindleyi Klotzsch ex Lindl. Syn. Conif. 59. 1847. C. Benthamii 
Endl. loc. cit. C. lusitanica var. Benthamii Carriere, Traite" Ge"n. 
Cbnif. ed. 2. 155. 1867. C. Knightiana Knight & Perry ex Gordon, 
Pinet. 61. 1858. C. lusitanica var. Knightiana Rehder, Journ. Arnold 
Arb. 1 : 52. 1919. C. Benthamii var. Knightiana Masters, Journ. Linn. 
Soc. 31: 340. 1896. Cipres; Tsicap (Jacaltenango) ; Tzis (Huehuete- 
nango, fide Tejada) ; Quisis (Quiche", fide Tejada) ; Chinchac; Paxaque; 
Ksis (Volcdn de Santa Maria, Quezaltenango). Figure 5. 

Native probably at about 2200-3300 meters, but widely planted 
and naturalized, and sometimes appearing native at considerably 
lower elevations; the following departments are those in which the 
tree probably is native: El Progreso (Sierra de las Minas); said to 
grow in Jalapa in region of La Soledad; Chimaltenango; Quiche" (?); 
Totonicapan; Quezaltenango; San Marcos; to be seen in cultivation 
or naturalized in all the departments. Central and southern Mexico. 



28 FIELDIANA: BOTANY, VOLUME 24 

A giant tree when well grown, sometimes 30 meters high or more, with a large 
or small, broad or columnar crown, the bark shredded and often separating in long 
narrow strips, reddish brown; branches in the common form spreading, often with 
somewhat refracted tips and pendulous branchlets, the smaller branchlets irregu- 
larly branched, slightly compressed; leaves closely imbricate, dull or dark green to 
glaucescent, 4-ranked, 1-2 mm. long, ovate, acute, appressed, with a dorsal glan- 
dular pit; cones pedunculate, globose, covered with a pale bloom, 12-15 mm. in 
diameter; scales 6-8, terminated by an elongate, pointed, and usually curved 
mucro; seeds yellowish. 

A few varieties or forms of this species have been described, but 
they are based either on herbarium specimens or on plants cultivated 
abroad, and apparently are of the very slightest importance and have 
no relationship with the rather obvious forms recognizable in the 
wild trees forms so distinct that they are given distinctive names 
by the Indians. Of the named forms one is C. lusitanica var. Ben- 
tkamii (Endl.) Carriere, described as a pyramidal tree, the branchlets 
regularly pinnate-branched and disposed in one plane, the smallest 
branchlets slightly compressed. C. lusitanica var. Knightiana 
(Knight & Perry) Rehder is said to be similar to var. Benthamii, 
but the branchlets are still more compressed and very regularly and 
evenly pinnate-branched. 

There are two conspicuous local variations that are of horticul- 
tural importance, at least in Guatemala, and they are easily recog- 
nizable in the field although not in herbarium specimens. The first 
of these is the local weeping cypress or "cipre"s lloron." In this the 
ultimate divisions of the branches, at least the lower ones, are very 
long and slender and laxly pendent, after the manner of the weeping 
willow (Salix babylonica), although on a much less exaggerated scale. 
We have not noted this form in the forest, but many trees are planted 
for ornament through the mountain regions, as, for example, about 
Antigua, where the trees are highly esteemed. 

The most conspicuous and distinctive variant of Cupressus lusi- 
tanica is the so-called cipres romano ("Roman cypress") for which 
the Occidente and particularly the city of Quezaltenango are noted. 
In this the crown of the tree is very long and narrow and of columnar 
form, looking exactly like the celebrated cypresses that dominate 
many Italian landscapes. In the wild state the columnar form seems 
to be confined to the mountains near Quezaltenango, and perhaps 
wholly to the Volcano of Santa Maria. On the upper slopes of this 
peak, in spite of the unfavorable loose sandy soil that has a marked 
tendency to slide to the foot of the mountain, cypress trees are plenti- 
ful, although not (where we have seen them) forming dense forests. 
Here the two forms, with broad and with columnar crowns, both 




FIG 5 Cupressus lusitanica. a, Cone (XI). 6, c, Seed (slightly enlarged). 
d, Habit of fruiting branch (X %). e, Habit of leafy branch (X \ft. 



29 



30 FIELDIANA: BOTANY, VOLUME 24 

grow, sometimes together and sometimes apart. Standing on some 
eminence near the base of the volcano one may look upward and map 
the distribution at a glance. It is believed that both the columnar 
and weeping cypresses come true from seed; at least this is the local 
belief. 

The native cypress forests of Guatemala are very fine and beauti- 
ful, but they are being none too gradually destroyed. The best stand 
of lumber is or was that on the Cerro de Tecpam in Chimaltenango, 
but this has been greatly depleted by lumbering operations, being on 
privately owned ground. The large trees left here are magnificent, 
the trunks very thick and tall, and often covered with great masses 
of soft green mosses as large as sofa cushions. The next best, and 
perhaps even more beautiful stands, are in San Marcos and adjacent 
Quezaltenango. Those along the road above Ostuncalco are espe- 
cially handsome the trees relatively young, but close together, and 
the trunks consequently very tall, slender, and straight. The cypress 
usually but not always is associated with Pinus Ayacahuite and Abies. 
Very few smaller plants are able to thrive in the denser forest, where 
but little light reaches the ground. In favorable places the trees seed 
profusely and seedlings are abundant. Many of the young plants 
doubtless are destroyed by the ever-present sheep. The senior author 
once was amazed when walking along the border of one of these mixed 
cypress-fir forests between San Francisco El Alto and Momostenango, 
to lift his eyes and see grazing there a small flock of llamas. It had 
the appearance of a hallucination, until it was remembered that the 
government of Peru some time previously had presented these ani- 
mals to Guatemala, and an attempt was being made to acclimatize 
them here. 

It is not possible, except by inference, at the present time to deter- 
mine the original distribution of cypress in Guatemala. The trees are 
seen in practically every settled part of the country, in the lowlands 
only as isolated trees in parks or about houses, but at 1300 meters or 
higher often in great groves that from a distance appear to be native, 
until closer inspection reveals that the trees are planted in rows. It 
may well have been that before any of the land was cleared, long 
before the Conquest, cypress covered most of the moister mountains 
above about 2200 meters, but this is purely guesswork. There can 
be no doubt that the forests were formerly much more extensive than 
today. This tree has been highly recommended by Wilson Popenoe 
and others for use in reforestation in tropical America. The prac- 
ticability of this is proved by the fact that in Guatemala it has been 
so used for probably several generations. The tree shows extraordi- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 31 

nary adaptability to climatic conditions, and in the parks it grows 
luxuriantly and rapidly, almost if not quite down to sea level. In 
recent years it has been much planted under government direction, 
in a laudable attempt to foster tree planting on all available land not 
used for ordinary crops. 

The native cypress is highly esteemed for ornamental planting, 
and there are numerous fine plantings of the trees in scattered places. 
One of the most imposing is the majestic avenue of great trees border- 
ing the entrance to San Marcos from the south, and other avenues 
perhaps quite as fine are seen about Chimaltenango. Perfect trees 
of the romano type may be seen about Antigua, as in front of the 
historic church of La Merced. The cypress makes excellent hedges 
and is much planted for this purpose. The hedges may be trimmed 
to any height and become very dense by continued pruning. About 
Quezaltenango and other cities there are striking examples of the 
topiarist's "art" the trees trimmed in the forms of peacocks and 
other animals. Among the historic trees of Guatemala is the famous 
tree, now dead but still standing in 1941 at least, at Conception Chi- 
quirichapa, near Quezaltenango, under which Pedro de Alvarado and 
his men are said to have rested before passing on to the conquest of 
what is now Quezaltenango. Its trunk is almost 12 meters in cir- 
cumference, but only the snagged stubs of a few of the old branches 
remain. 

Cypress lumber is highly esteemed in Guatemala for construction 
purposes, and the trees still are being cut. It is stated that the wood 
is used commonly for the sounding boards of guitars and mandolins. 
The young branches are used in great quantities for decorations dur- 
ing all the fiestas, and they are employed in large amounts for making 
coronas or wreaths for cemeteries. While the pines of Guatemala 
have been reduced by fungus and insect pests, the cypress seems 
resistant to such enemies. In some places the branches are much 
deformed by a rust that produces growths similar to cedar apples, 
but trees so infested seem to suffer but little damage. 

The specific name liLsitanica ("Portuguese") for this tree is an 
unfortunate and misleading one. Seeds were supposedly taken from 
Mexico by Spanish friars, and reached Portugal, where the tree was 
grown and dispersed to other parts of Europe, receiving finally the 
name Cupressus lusitanica because it was believed to have originated 
in Portugal. Professor Martinez, in his work on "Los Cupressus de 
Mexico" (Anal. Inst. Biol. Mex. 18: 71-149. 1947) treats the mate- 
rial from Guatemala under the name of C. Lindleyi Klotzsch, apply- 
ing that name likewise to a great mass of Mexican material, at the 



32 FIELDIANA: BOTANY, VOLUME 24 

same time differentiating C. Benthamii Endl. as a less commonly dis- 
tributed Mexican species confined to the states of Hidalgo, Veracruz, 
and Puebla. Since he cannot match any material originating from 
Mexico with that sent by Dr. Melo de Figueiredo from what is con- 
sidered to be the first tree of C. lusitanica cultivated in Bussaco, 
Portugal, he concludes that C. lusitanica cannot be native to Mexico 
(loc. cit. 89-90) . Moreover, he retains C. lusitanica as distinct from 
both C. Lindleyi and C. Benthamii, distinguishing them by what he 
considers to be good differences (see pp. 98 and 104). Inasmuch as 
there exists considerable variation in wild and cultivated specimens 
of Guatemalan Cupressus and in what would pass as similar or iden- 
tical material from Mexico, not only in shape of tree, position and 
thickness of branches, size of fruits, but also in other details, it is not 
surprising that various authors have reached different conclusions 
regarding the disposition of the variations encountered. The differ- 
ences mentioned by Martinez (loc. cit.) for separating these three 
names do not appear justified in our opinion, as there is considerable 
variation and overlapping of characters used. As C. lusitanica is the 
oldest name used for what we consider to represent a complex taxon 
present in Guatemala and Mexico, it seems advisable, as some other 
authors have done, to adopt its usage as the conservative course of 
procedure, instead of disregarding it completely. 

JUNIPERUS L. Red Cedar; Jumper 

Evergreen aromatic trees or shrubs, sometimes low or even prostrate, most 
often with thin shredded bark, the branchlets spreading, ascending, or irregular, 
the secondary branchlets produced on all sides of the primary ones; leaves small, 
opposite or ternate, either acicular and somewhat spreading or scale-like and 
appressed, the needle-like leaves in whorls of 4, with bluish or whitish lines on the 
upper surface, the scale-like leaves in 3-4 rows, both kinds of leaves often found on 
the same plant at different stages of development; flowers dioecious or monoecious; 
staminate catkins small, ovoid or oblong, terminal or in the leaf axils, the flowers 
yellow, consisting of numerous, opposite or ternate, 1-celled anthers; pistillate 
flowers minute, globose, greenish, composed of 3-8-pointed fleshy scales or bracts, 
some or all of these bearing 1-2 ovules within, the fruit finally becoming fleshy and 
berry-like, with a strong resinous odor, usually maturing the second year but some- 
times the first or third year; seeds generally 1-6, usually ovoid, terete or angulate, 
often sulcate, brown or buff, with a conspicuous hilum at the base. 

About 40 species, in the northern hemisphere, chiefly in temper- 
ate regions, in the tropics usually confined to the mountains. The 
following are the southernmost American species. The wood in this 
genus is brown to reddish or purplish, with a strong and pleasant 
fragrance; very fine and of uniform texture, firm and fairly hard, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 33 

easy to cut in any direction, holds its place well when manufactured, 
and is highly resistant to decay. In the United States cedar wood is 
used principally for making lead pencils, for which purpose no satis- 
factory substitute has been found. It is utilized also for making 
chests in which woollen goods are stored, since it is repellent to clothes 
moths and other insects. Little or no use is made of the cedar wood 
in Guatemala except for firewood. The volatile oil of the berries dis- 
tilled with spirits is a beverage, gin, in Spanish ginebra, both words 
derived from the geographic name Geneva (Ginebra in Spanish) . A 
decoction of the leaves of various species has been used as a teniafuge 
and abortifacient, but its use is dangerous. The bark is rich in tannin 
and is sometimes utilized for tanning leather. 

Adult scale-like leaves acute, mucronate, 1-1.3 mm. long; superficial epidermal 
cells very fine and numerous under a microscope; margins of the scale-like 
leaves conspicuously erose-denticulate; fruit 1-seeded or rarely 2-seeded; seeds 
large, 8 mm. long, firmly attached to the pericarp J. comitana 

Adult scale-like leaves cucullate-rounded or obtuse at the apex, not mucronate; 
superficial epidermal cells relatively larger and fewer when seen under a micro- 
scope; margins of the leaves minutely erose to subentire; seeds only 4-5 mm. 
long, easily separating from the pericarp J. Standleyi 

Juniperus comitana Martinez, Anal. Inst. Biol. Mexico 15: 12. 
/. 5-8. 1944. Sicop, Cipres (Baja Verapaz). 

Mostly on rather dry and open, rocky hillsides, 1200-2150 meters; 
Baja Verapaz (region of Santa Rosa) ; Zacapa (Sierra de las Minas) ; 
Huehuetenango. Chiapas (the type from Comitan). 

A large shrub or a tree, seldom more than 12 meters high, the bark shredded, 
separating into tough, reddish brown strips or long scales; branchlets slender, 
4-sided, separated from one another, 0.8-1 mm. thick, the ultimate ones 1.5-2 cm. 
long; adult leaves scale-like, ovate, acute, mucronate, bright green, 1-1.3 mm. long, 
dorsally depressed at the middle and glandular or eglandular, the margins conspic- 
uously erose-denticulate, the apex and margins paler than the rest of the leaf; 
needle-like leaves subulate or linear-subulate, 5-11 mm. long, 0.8-1 mm. wide; 
fruit containing 1 or rarely 2 seeds, bluish, glaucous, with thin sweet resinous 
flesh, 9-10 mm. long, 5-8 mm. broad. 

This tree is very local in Guatemala. In general appearance as 
well as in habit of growth it is conspicuously different from J. tetra- 
gona, usually occurring as an isolated tree with rather open crown. 
Even the smaller plants are strictly erect. Indians questioned in 
Baja Verapaz insisted that the tree was the common Cupressus of 
Guatemala, but the fruits of Juniperus and Cupressus are quite unlike. 

Juniperus Standleyi Steyermark in Standl. & Steyerm. Field 
Mus. Bot. 23: 3. 1943 (type from upper slopes of Volcan de Tacana, 



34 FIELDIANA: BOTANY, VOLUME 24 

San Marcos, Steyermark 36137). Huitun (Huehuetenango; accent 
on the first syllable) ; Huitd (Huehuetenango) ; Cipres. Figure 6. 

High mountains, often on limestone, 3000^4100 meters; Huehue- 
tenango (Sierra de los Cuchumatanes; mostly at high elevations, but 
sometimes at lower altitudes) ; San Marcos (near summit of Volcan de 
Tacana). Chiapas (Volcan Tacana). 

Plants sometimes prostrate and forming mats only 30 cm. high, or often low 
broad spreading shrubs about a meter high, becoming trees as much as 15 meters 
high and then often forming dense forests; branches reddish brown, the ultimate 
branchlets with scale-like leaves often excessively numerous, dense, and crowded, 
mostly 1.1-1.4 mm. thick and 5-20 mm. long; adult leaves scale-like, ovate or 
rounded-ovate, rounded or obtuse and somewhat cucullate at the apex, deep green, 
1.5-1.7 mm. long, depressed at the middle, glandular dorsally, the margins thick, 
incurved, minutely erose or subentire, the tip and upper margins the same color as 
the rest of the leaf; needle-like leaves lance-subulate, acuminate, 4-5 mm. long, 
1.5-1.7 mm. wide at the base; fruit containing 3-4 seeds, bluish green or bluish, 
7-9 mm. in diameter, solitary on a short curved pendulous pedicel. 

This cedar is found most abundantly and almost exclusively at 
high elevations, 3000 meters or higher. On the Volcan de Tacana it 
is confined to moist ravines or shaded rocky slopes toward the sum- 
mit, where most of the plants are prostrate or nearly so. In the 
Cuchumatanes the tree is abundant, and perhaps the most distinctive 
tree of high elevations, forming there small or large groves of very 
dense growth, frequently on rough limestone slopes where there is 
but little soil. These forests are so dense and there are so many dead 
branches on the ground that frequently it is difficult to make one's 
way through them. Perhaps because of the cold and fog prevailing 
in these places, the forests have a forbidding aspect which is enhanced 
by the many dead gnarled trees with jagged whitish branches stripped 
of bark, the many gray lichens clinging to the branches, and the 
numerous ravens that perch in the trees, croaking as if in pain. In 
such dense forests the little other vegetation consists chiefly of strag- 
gling shrubs of Mahonia, Holodiscus, and Rubus trilobus. Most of 
the cedar trees in the Cuchumatanes are gnarled, and few are of 
symmetrical growth. They vary in shape but most have low, broad, 
rather depressed crowns. The branches often have conspicuous knots 
and also long fusiform swellings as thick as a man's arm, doubtless 
caused by some Gymnosporangium or similar rust. In open places 
about the denser forest the cedar often grows as a low, depressed and 
spreading but not prostrate shrub, a meter high or somewhat lower. 
The trees were in bloom in the Cuchumatanes at the end of Decem- 
ber, shedding clouds of pollen when the branches were shaken. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 35 




FIG. 6. Juniperus Standleyi. a, Seed, ventral view (X 5). b, Seed, lateral 
view (X 5). c, Pistillate flowers on tip of branchlet (X 2). d, Staminate catkins 
on end of branchlet ( X 3). e, Habit of fruiting branch (X Yd- 



THUJA L. Arbor-vitae 

Evergreen aromatic monoecious trees or shrubs with spreading, erect, or 
pendent branches, the crown usually pyramidal, the bark thin, irregularly scaly; 
branchlets compressed, disposed in one plane, pinnately branched, the smaller ones 
deciduous; leaves on young plants needle-like and spirally arranged, the normal 
adult leaves small, scale-like, imbricate in 4 rows, decussate, appressed, sometimes 
dorsally glandular; flowers small, terminal, on short branchlets; staminate flowers 
ovoid, yellow, the 6-12 decussate stamens each with 2-4 anther cells; pistillate 
flowers consisting of 8-12 scales in opposite pairs, only the middle or lower scales 
fertile, each of the fertile scales with basilar ovules within; cones globose-ovoid to 
oval-oblong, the scales with a thickened ridge or umbo at the apex; seeds 2 under 
each scale, thin, with broad lateral wings, or thick and wingless; cotyledons 2. 

Six species, 2 in North America, 4 in eastern Asia, in temperate 
regions. 



36 FIELDIANA: BOTANY, VOLUME 24 

Thuja orientalis L. Sp. PL 1002. 1753. Cipres; Incienso. 
Native of eastern Asia, often planted for ornament in Guatemala. 

A shrub or a small symmetrical tree, the branches slender, ascending or pend- 
ent, the bark reddish brown; leaves of the main axis with a free, somewhat spread- 
ing apex, those of the lateral branchlets closely appressed, rhombic-ovate, acute, 
bright green, with a narrow linear-elliptic gland in the center dorsally; cones 
globose-ovoid, 1.2-2.5 cm. long, somewhat fleshy and bluish before maturity; scales 
usually 6, ovate, the uppermost pair sterile, each with a horn-like mucro near the 
apex; seeds ovoid, brown. 

Most of the plants seen in Guatemala are mere shrubs, but some- 
times small trees may be found. They resemble the cypress but are 
distinguished by their flattened branchlets. Some of the named vari- 
eties and forms are represented locally, distinguished by slender 
drooping branches, golden foliage, and other characters. 

PINACEAE. Pine Family 

Usually large trees, the leaves persistent, resinous, spirally arranged, solitary 
or fasciculate; flowers naked, usually monoecious, subtended by scales; stamens 
numerous, the sporangia 2, on the lower side of the scale; pistillate flowers with 
numerous, spirally arranged scales, these free or sometimes connate; fruit a cone, 
usually woody, closed until maturity or later, composed of the much thickened 
and indurate scales; seeds usually winged along one margin; embryo with several 
cotyledons. 

Nine genera are known, the rather numerous species almost all in 
the northern hemisphere. Only the following genera are native in 
Central America but four others are native in North America, and 
three of them, Pseudotsuga, Tsuga, and Picea, extend into Mexico. 
Besides the genera discussed below, a species of Larix, perhaps the 
European L. decidua Miller, has been observed in cultivation in 
Guatemala City. The species of Larix (in English, larch) resemble 
the pines in their leaves, but the leaves are soft and deciduous. There 
is in cultivation also in Guatemala City, in the Parque Central and 
probably elsewhere, the Norway spruce, Picea Abies (L.) Karsten, 
native of northern and central Europe, a tree highly esteemed for 
ornamental purposes in the United States. 

Leaves solitary, not subtended by a basal sheath; scales separating from the axis 
of the cone at maturity Abies 

Leaves fasciculate, the fascicle subtended by a sheath; cone scales persistent on 
the axis at maturity Pinus 

ABIES Miller. Fir 

Reference: Alfred Rehder, The firs of Mexico and Guatemala, 
Journ. Arnold Arb. 20: 281-287. 1939. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 37 

Usually large, evergreen trees, symmetrical in growth, the crown pyramidal or 
conic-oblong, the branches spreading, whorled; trunk straight, the bark smooth 
and thin on younger trees, in age becoming thick and furrowed at the base; winter 
buds with or without resin; leaves spirally arranged but because of a basal twist 
usually spreading and 2-ranked, persistent on dried branches, linear, flat and sul- 
cate above, beneath usually with 2 white stomatiferous bands, sessile, contracted 
above the base, leaving a circular scar when fallen, with 2 (in Guatemalan species) 
or rarely 4-8, marginal or median resin canals, and with usually 2 vascular bundles; 
aments axillary, appearing in early spring from buds formed the previous season 
on branchlets of the year, surrounded by involucres of the enlarged scales of the 
flower buds; staminate catkins pendent on branches above the middle of the tree, 
oval to cylindric-oblong, the anthers yellow or scarlet; pistillate catkins globose, 
ovoid, or oblong, erect on the uppermost branches, the scales numerous, 2-ovulate, 
imbricate; cones erect, ovoid to oblong-cylindric; cone scales closely imbricate, thin 
and coriaceous, incurved and broad or truncate at the apex, narrowed at the base 
into a long stipe and subtended by narrow, exserted or enclosed bracts, the scales 
falling at maturity from the persistent axis; seeds ovoid or oblong, with large thin 
wings; cotyledons 4-10. 

About 40 species in temperate regions of the northern hemisphere, 
in America ranging southward to Guatemala, in the Old World to 
northern Africa and the Himalayas. The genus includes some of the 
stateliest and handsomest of all the conifers. The wood is almost 
colorless to brown with a reddish or pinkish tinge, of medium luster, 
very light, soft, and weak to moderately heavy, hard, and strong. 

Abies guatemalensis Rehder, Journ. Arnold Arb. 20: 285. /. 1, 
j-m. 1939. A. tacanensis Lundell, Amer. Midi. Nat. 23: 175. 1940 
(type from Volcan de Tacana, Chiapas); A. guatemalensis var. taca- 
nensis (Lundell) Martinez, Anal. Inst. Biol. Mexico 19: 70-73. 
/. 51-52. 1948. Pinabete. Figure 7. 

Moist or wet forests of the high mountains, mostly at 2700-3500 
meters; Quich (Cerro Maria Tecun) ; Totonicapan (Cumbre del Aire; 
region of Desconsuelo and between San Francisco El Alto and Mom- 
ostenango) ; Huehuetenango (type from Cumbre del Aire, J. H. Fault 
13104, perhaps rather in Totonicapan; Sierra de los Cuchumatanes, 
at Chancol, region of San Mateo Ixtatan, and elsewhere) ; Quezalte- 
nango (Volcan de Zunil; mountains southeast of Palestina); San 
Marcos (volcanoes of Tajumulco and Tacana; region of Serchil). 
Chiapas (Volcan de Tacana), Oaxaca, and Guerrero. 

A tree as much as 45 meters high with a trunk almost a meter in diameter, or 
perhaps even larger, the branches dark or grayish brown, the young twigs rufous- 
brown or castaneous, sparsely hirtellous near the apex; leaves appearing 2-ranked, 
spreading-ascending or almost divaricate, linear, 1-4.5 cm. long, 1-2 mm. wide, 
obtuse and usually emarginate at the apex, lustrous and dark or rather light green 
above, usually silvery beneath, the upper surface sulcate for all or most of its 



38 FIELDIANA: BOTANY, VOLUME 24 

length, the costa elevated beneath, the margins recurved, the stomata conspicuous 
beneath; resin canals 2, subepidermal, the hypoderm well developed, interrupted; 
fibrovascular bundles 2, approximate but distinct; cones subsessile, 8.5-11.5 cm. 
long, 4.5-5 cm. in diameter; bracts cuneate-obovate or oblanceolate, half as long 
as the scales or somewhat exceeding them; broadly rounded or truncate and erose- 
denticulate at the apex, the cusp usually exserted; scales broadly cuneate-obovate 
or transverse-oblong, 2.7-3 cm. wide, 1.5-2.2 cm. high, the margins hirtellous- 
puberulent outside; seed cuneate-obovoid, 8-10 mm. long, pale brown, the wings 
obovate, 1-1.5 cm. long, 1.4-1.5 mm. wide. 

Until recently the Guatemalan fir was confused with the Mexican 
A. religiosa (HBK.) Schlecht. & Cham., which received its specific 
name because the branches were used as decorations in churches. 
A. tacanensis of Volcan de Tacana was supposed to be distinguished 
from A. guatemalensis by its narrower, more crowded leaves with 
margins rounded in cross section, the upper leaf surface sulcate only 
in the lower two-thirds, and the longer bracts of the cones. Martinez 
(Anal. Inst. Biol. Mexico 19: 70-73. 1948) has attempted to retain 
A. tacanensis as a variety of A. guatemalensis on the basis of leaf, 
cone, and scale measurements, differences in apex of leaves and wings 
of the seeds, and internal structure of the leaves. However, the am- 
ple Guatemalan material now available varies in all these characters, 
and many of the variations may be found on the same tree. 

The fir is one of the outstanding trees of Guatemala, although of 
local distribution, dependent in large part upon its altitudinal re- 
quirements. It is abundant in some localities, where it forms dense 
and handsome forests of limited extent, usually in association with 
Pinus Ayacahuite and Cupressus. Occasionally, as at the type local- 
ity, the trees are spaced, and sometimes isolated. It is somewhat 
surprising that the tree was not practically exterminated long ago, 
and its persistence at present is largely a result of the fact that most 
of the best stands are on national lands, where their cutting is pro- 
hibited. The best forests are so dense that little vegetation is found 
on the ground beneath the trees only a few low shrubs of rather 
weak growth, mosses, and a limited number of herbs. The trees are 
very rarely planted in parks and fincas, as in the hotel patio at Solola. 
Some are said to be planted in the region of Coban but we have not 
seen them, and the climate there probably would not agree with 
them. In their native habitat the trees do not reproduce very freely, 
but some young plants are found. The isolated trees at Cumbre del 
Aire have been found repeatedly loaded with cones, but the constant 
grazing by sheep probably destroys the seedlings. 

Because of its scarcity, no lumber of this tree is available in Gua- 
temala, except on rare occasions. It is stated, however, that the wood 








FIG. 7. Abies guatemalensis. a, Apex of leafy branch (X 1). b, Portion of 
branch with staminate catkins (XI). c, Cone scale, inner side (XI). d, Seed 
(X1 1 A). e, Needle, lower side (X 1)4). f, Cone (XI). 



39 



40 FIELDIANA: BOTANY, VOLUME 24 

is in demand for making the hand looms on which the beautiful 
Indian textiles are woven. The branches often are cut in the moun- 
tains to make ramadas or temporary shelters. The principal use of 
the tree, however, is for decorative purposes. The branches are much 
used, when available, as decorations in churches and dwellings, and 
substantial quantities of them are available, probably cut surrepti- 
tiously on government land. Since this is the only native tree that 
affords Christmas trees similar to the best ones of Europe and the 
United States, the small trees are much in demand for this purpose 
among the foreign residents. Fine Christmas trees were observed on 
sale in Huehuetenango for as little as twelve cents, while in Guate- 
mala, to which they must be carried for long distances, they usually 
cost five dollars or more. 

PINUS L. Pine 

References: G. R. Shaw, The pines of Mexico, pp. 1-29, pis. 1-22. 
1909; The genus Pinus, Publ. Arnold Arb. 5, pp. 1-96. 1914. M. Mar- 
tinez, Las pinaceas mexicanas, Anal. Inst. Biol. Mexico, 345 pp. 1945; 
Los pinos mexicanos, ed. 2. 1948. E. E. M. Loock, The pines of Mex- 
ico and British Honduras, in Union of South Africa Dept. For. Bull. 
35. 244 pp. 1950. 

Evergreen trees or rarely shrubs, the branches whorled, spreading, the bark 
furrowed, broken into plates or sometimes rather smooth; buds covered with imbri- 
cate, more or less resinous scales; leaves and branchlets of two kinds, the incon- 
spicuous primary leaves arranged on long shoots and usually reduced to small 
scarious bracts, the conspicuous green secondary leaves arising in the axils of the 
primary leaves or bracts on short undeveloped shoots or branches; leaves in clus- 
ters of 2-6 (rarely solitary), semiterete or trigonous in cross section, much elongate, 
slender, needle-like, the cluster surrounded at the base by a thin colored sheath 
composed of 8-12 bud scales, the sheath often deciduous; flowers monoecious, in 
yellow, orange, or scarlet, catkin-like clusters at the ends of the old shoots or at 
the bases of the young ones, composed of many spirally arranged, 2-celled anthers; 
pistillate flowers lateral or subterminal, in short, dense, greenish or purplish spikes, 
these composed of numerous, spirally arranged, 2-ovulate scales, each scale sub- 
tended by a small bract; cones maturing at the end of the second or third season, 
subglobose to long-cylindric, symmetric or asymmetric, the woody scales appressed 
and closed before maturity; apex of the scale usually much thickened, the exposed 
part (apophysis) generally rhombic in outline, transversely carinate and usually 
with a prominent umbo, this sometimes ending in a spine or prickle; seeds small or 
large and nut-like, with or without a wing; cotyledons 4-15. 

About 65 species, all in the northern hemisphere except for one 
or more that extend into the Malayan Archipelago. Only the follow- 
ing are found in Central America, but many others are native in 
Mexico and the United States. Most of the tropical pines are con- 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 41 

fined to mountain regions, but sometimes they descend to sea level. 
Several exotic pines are planted for ornament in Guatemala, one of 
them being (in the Jardin Botanico at least) Pinus halepensis Miller, 
native of southern Europe and western Asia, the Aleppo pine. 

Pine trees are of the highest economic importance, being the most 
important source of lumber and resin of all trees. In Guatemala 
they are also the most important single source of lumber, and 
are used for almost every purpose for which wood can be utilized. 
Some of the lumber is sawed in sawmills, and much is sawed by hand. 
In the latter case the log usually is placed over a shallow pit, and two 
men, one above and one below, manipulate a cross-cut saw, painfully 
working out large boards. One must admire the skill of these work- 
men, who obtain wide boards almost as uniform in thickness as those 
obtained by more modern processes. The rough boards often are 
transported for long distances up and down the mountains upon 
men's backs, or more often dragged along the ground, a task in which 
women and girls sometimes engage, as we have seen them in the 
mountains between Totonicapan and Quezaltenango. 

Pine wood also is an important firewood in Guatemala, although 
not a very good one. It is used generally only when better fuel is not 
available. In the rural regions, especially among the Indians, a pine 
torch is often used for illumination along the roads and trails or 
even in the houses, where its cheapness is an important factor in its 
use. Small billets of fat pine or ocote are sold in all the small supply 
shops for lighting fires, especially charcoal; it is amusing to watch 
women and children, before they make a purchase, inspecting each 
item of the stock with almost as much care as if they were buying a 
new garment. Some resin and turpentine are doubtless gathered 
in Guatemala, but we have no reliable information regarding this 
subject. 

The two most abundant and characteristic trees of Guatemala as 
a whole are pines and oaks, and these often form mixed forests. Fre- 
quently, however, the pines form beautiful pure stands of great 
extent. One of the pines, P. caribaea, is isolated in the lowlands of 
the Atlantic coast. Of the white pines, P. Ayacahuite is found only 
at high elevations, usually associated with Abies and Cupressus, 
whereas the southern variety of the white pine of the eastern United 
States, P. Strobus var. chiapensis, occurs in moist forests at lower 
and middle elevations of the Sierra de los Cuchumatanes in the de- 
partments of Quiche" and Huehuetenango. The other three are much 
alike in their characters, widely distributed at middle elevations or 
sometimes at rather high ones. Taken together they form a distinc- 



42 FIELDIANA: BOTANY, VOLUME 24 

tive group, all three species much alike in their habits and habitats, 
and not easily separable in either the field or the herbarium. These 
three cover or formerly covered most of Guatemala lying at middle 
elevations, except in the more humid areas like the Pacific slopes of 
the volcanoes, where a mixed forest of broad-leafed trees prevails. 
There are still very extensive pine forests in Guatemala, some of them 
much over-grazed by cattle and sheep, and much of them consisting 
only of small trees. Where transportation is poorer than usual, hand- 
some stands of large trees may be found, as in the mountains of Baja 
Verapaz. A vast part of Guatemala that once must have been cov- 
ered with pine forest is now under cultivation, and some of it has 
been for centuries. At the present time the pine forests are suffering 
severely not only from inroads made by man but even more from 
the attacks of insect and fungus pests. In almost every part of Gua- 
temala one sees hundreds of acres or many square miles of land where 
practically all the pine trees are dead but still standing. It is impos- 
sible to believe that the disease situation always has been so bad as 
at present, for if it had been, the forests certainly would have been 
destroyed long ago. These pests have been studied at least super- 
ficially, and have been found to be numerous various kinds of fungi 
and several insects of different groups. The most striking disease 
attacking the pines is the so-called big-cone disease, which is well 
known also in Mexico. In Guatemala this is caused by the larvae of 
a moth. Pine cones in which the larvae develop are malformed and 
attain several times their normal size. About Coban after a heavy 
rain one sees pine trees (Pinus oocarpa) loaded with golden-yellow 
cones, as if they had been freshly gilded. This appearance is caused 
by the great masses of spores of a rust fungus. 

Especially in the central region one frequently sees pine trees that 
are abnormal in appearance, the tall slender trunks naked except at 
the top, where there is only a small tassel-like crown. The branches 
have been cut for use as decorations for fiestas and celebrations. On 
festive occasions it is customary, a custom doubtless dating back to 
preconquest times, to cover the floors and even sometimes the streets 
with fresh green pine needles. More than that, the fresh branches 
are used for adorning walls of houses, outside and inside, for decorat- 
ing altars, and for making the characteristic arches across roads and 
streets. Pine branches are a ceremonial offering by the mountain 
Indians to roadside crosses and shrines, and Sapper believes that they 
must have been offered likewise to the ancient pagan gods. He states 
that in parts of Alta Verapaz where pine trees do not grow naturally 
they are planted, so that branches may be available for this purpose. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 43 

A more prosaic use for pine needles is their addition to adobe bricks, 
where they serve the same purpose as cereal straw. 

Pine trees often are planted for ornament or with an eye to their 
use for lumber in almost all the mountain regions of Guatemala. The 
cutting of seedlings or small trees is punishable by a severe fine, a 
punishment that has often been inflicted in recent times. The term 
ocote, which is applied to fat pine slivers, is derived from the Nahuatl 
ocotl, the Nahuatl term for pine tree. A pine of the Lacandon region, 
probably P. caribaea, is said to be called "tote" by the Lacandones. 
Other Indian names of Guatemala applied to pine trees, probably 
to all species without discrimination are "chaj" (Huehuetenango) ; 
"titzlum" (Huehuetenango); "chaj" (Totonicapan). The following 
names are used in the Guatemalan languages indicated: "chaaj" 
(QuichS) ; "tax," "caxta" (Jacalteca) ; "taj" (Maya of Pete*n) ; "tajte"" 
(Tzental); "sacal taj" (Choi); "tiatie"" (Quecchi); "chaj" (Pocom- 
chi); "tza" (Aguateca); "tzaj" (Mame); "teccamanil" (San Marcos). 
"Chajul" is a Quich word signifying a pine billet used as a torch. 

Some of the Mexican pines produce edible seeds or pinones that 
are very good to eat, especially when roasted. It is said that the 
Mexican nut pines, presumably Pinus cembroides Zucc., have been 
planted at a few places in the Occidente of Guatemala. Pinones im- 
ported from Mexico are offered for sale here, but at high prices. 

Leaf sheaths deciduous; mature leaves silvery or whitish on one side; cone scales 
with the umbo terminal and unarmed; bark of mature trees relatively smooth 
or slightly furrowed. 

Apophysis of cone scale elongated and reflexed; cones 15 cm. or more long, 
usually 3-5 times as long as broad; resin ducts 2-6; at elevations of 2000- 

4000 meters, widely distributed in Guatemala P. Ayacahuite 

Apophysis of cone scale not elongated, broadly rounded at apex; cones less than 
15 cm. long, usually 2-2 1/6 times as long as broad; resin ducts usually 3; 
at elevations of 800-1800, rarely 2000 meters, rare in departments of 

Quichd and Huehuetenango P. Strobus var. chiapensis 

Leaf sheaths persistent in age; mature leaves green on all sides; cone scales with 
the umbo dorsal, generally with a prickle or sharp-pointed process; bark of 
mature trees strongly furrowed or broken into plates. 

Leaves in fascicles usually of 3; young shoots multinodal; trees of low elevations, 
at 600 meters or lower; resin ducts of the leaves internal, i.e., touching the 
endoderm P. caribaea 

Leaves in fascicles usually of 5-6, sometimes in 4's; trees of the higher moun- 
tains, usually at 1500 meters or higher; resin ducts of the leaves medial or 
septal. 

Branches of young trees and upper part of stem smooth, not covered with 
rough scales, the base of the scales not prominent and more or less sub- 
merged in the wood; leaves slender, 0.6-1 mm. broad, often pendent, 
curving, or laxly spreading; resin ducts medial, i.e., not touching the 
hypoderm or endoderm P. pseudostrobus 



44 FIELDIANA: BOTANY, VOLUME 24 

Branches of young trees and upper part of stem rough, covered with rough or 
persistent scales or their prominent bases; leaves stout, stiff er and more 
rigid, 1-1.5 mm. broad, erect; resin ducts septal or medial. 
Cones borne on peduncles as much as 3.5 cm. long, long-persistent and late 
in opening, usually broadly ovoid or ovoid-conic, generally only 1.2-1.7 
times as long as broad; resin ducts septal, i.e., touching both the hypo- 
derm and endoderm P. oocarpa 

Cones sessile, opening at maturity and deciduous, conic-oblong to conic, 
usually 2-3 (sometimes 5) times as long as broad; resin ducts medial, 
i.e., not touching the hypoderm or endoderm. 
Trees with relatively long leaves and cones; leaves 15-45 cm. long. 

P. Montezumae 
Trees with relatively short leaves and cones; leaves 7-15 cm. long. 

P. Montezumae var. rudis 

Pinus Ayacahuite Ehrenberg, Linnaea 12: 492. 1838. Pino; 
Pino dulce; Pachd, (Volcan de Santa Maria, Quezaltenango). Fig- 
ure 8. 

Usually in moist mountains at high elevations, generally associ- 
ated with Abies or Cupressus or both, and often with broad-leafed 
trees, 2000-3300 meters or perhaps even higher; El Progreso (Sierra 
de las Minas) ; Zacapa (Sierra de las Minas) ; Jalapa (Montana Mira- 
mundo); Guatemala (Volcan de Pacaya); Quiche" (Cerro Maria 
Tecun); Huehuetenango; Totonicapan; Quezaltenango; San Mar- 
cos. Widely distributed in Mexico. 

A large tree, sometimes 50 meters high with a trunk almost 2 meters in diam- 
eter, the trunk often very tall and clean, the bark grayish white and smooth on 
young trees, on older trees becoming pale brownish or copper-brown and broken 
into shallow 4-angulate thin plates; young shoots uninodal; youngest branchlets 
slender, 4-8 mm. thick, rather smooth, grayish or light brown; leaves 5 in a fascicle, 
pale or grayish green above, silvery or whitish beneath, 5.5-22 cm. long, 0.6-1 mm. 
thick, flexible, somewhat serrulate, the stomata ventral only, the resin ducts exter- 
nal, the nbro-vascular bundle only 1; leaf sheaths soon deciduous; cones long-cylin- 
dric, gradually narrowed to the apex, pendent from long stalks, 22-45 cm. long, 
4-10 cm. thick, yellowish or reddish brown, dull to somewhat lustrous; cone scales 
prolonged, spreading and reflexed in age, relatively thin and flattened, the umbo 
terminal; seeds gray-brown, mottled with dark brown, with a well-developed wing 
3-3.7 cm. long and 8 mm. wide. 

This is the handsomest and most attractive of the pines of Guate- 
mala, often attaining a great height at high elevations, with a very 
tall, clean trunk frequently a meter or more in diameter. In some 
places, such as the slopes above Fuentes Georginas in Quezaltenango, 
it grows in the rather dense forest of broad-leafed trees, where it is 
conspicuous because of its unique foliage that glistens in the sunlight. 
This is one of the easily recognized Guatemalan pines, because of the 
deciduous leaf sheaths, and especially on account of the unique cones, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 45 




5. H. Grove 

FIG. 8. Pinus Ayacahuite. a, Leafy branch (X YZ). b, Cone (X H)- 



long and narrow with recurved cone scales and of very handsome 
appearance. 

Pinus caribaea Morelet, Rev. Hort. Cote d'Or 1: 105. 1851. 
P. hondurensis Loock, Union of South Africa Dept. For. Bull. 35: 
205-214. 1950, homonym of P. hondurensis Sneclauze, Conif. 126. 
1867. Pino; Pino bianco; Pino Colorado; Pino de ocote; Ocote; Sachaj 
(Alta Verapaz). Figure 9. 

Abundant on hillsides and plains at low elevations, 600 meters or 
lower; eastern Pete"n; Alta Verapaz (Savanna Sachaj, between Sachaj 



46 FIELDIANA: BOTANY, VOLUME 24 

and Sacacac) ; Izabal (lowlands, and on the slopes of Sierra del Mico, 
at about 600 meters or lower). Southern Quintana Roo; British 
Honduras; Atlantic lowlands of Honduras and Nicaragua south to 

the region of Bluefields; West Indies. 

- 

A rather slender tree, commonly 15-30 meters high with a trunk sometimes 
almost a meter in diameter, the bark gray, brownish-gray, or reddish-brown, broken 
into shallow longitudinal plates; buds pale chestnut or reddish-brown; branches 
horizontally spreading, the youngest ones 6-10 mm. thick, rather rough, dark or 
brownish-gray; leaves usually 3 in a fascicle, sometimes 4-5, rarely 2, grass-green 
on all sides, 5-25 cm. long, 1-1.5 mm. thick, rather stiff, serrulate, with stomata on 
all sides; resin ducts internal, mostly 3-4, the fibro-vascular bundles of the leaf 2, 
the sheaths persistent; cones oval or oblong-ovoid, reflexed and dehiscent at ma- 
turity, 4-12 cm. long, 3.5-6 cm. thick, dark or rufous brown, lustrous, on stalks 
1-1.5 cm. long; cone scales thickened at the ends, the umbo prominent and some- 
what mucronate; seed usually 4-5 mm. long, the wing of the seed usually remaining 
attached, its base thin or slightly thickened, 1.8-2 cm. long and 6 mm. wide. 

This is one of the easily recognized pines of Guatemala because 
of its limited range, which is widely separated from those of all other 
local pines. It has the distinction of extending farther southward 
than any other American pine, although Pinus oocarpa reaches al- 
most as far south, but in the interior of Nicaragua. The Caribbean 
pine is naturally one of the conspicuous trees of northeastern Guate- 
mala, and it is abundant in many places, forming extensive areas of 
pine savanna, similar to those of southern Florida (which are formed 
by a related species), and also extending well up on the slopes of 
Sierra del Mico in Izabal, where it forms open and rather sparse 
stands, sometimes in association with the cohune palm, and with an 
abundant undergrowth of small shrubs, many grasses and sedges, and 
a variety of small herbs, most of them of the same genera as those 
in the pine forests of the Florida Everglades. Riding on the train 
from Puerto Barrios toward Guatemala, one sees good examples of 
pine forest on the mountains of the Gualan region, and even better 
ones toward the coast, where in some places the train passes through 
pine savannas, in the region between Cristina and Virginia. In Gua- 
temala pines do not reach the coast, as they do at some places in 
Nicaragua. In British Honduras the Caribbean pine is even more 
abundant. It ascends to about 900 meters in the mountains and is 
said to occupy about one-third the area of the colony. The wood is 
of great importance locally for construction purposes. Some of 
the British Honduras trees are reported to attain a height of 30 
meters. 

The Maya name of this pine, in Quintana Roo and British Hon- 
duras, is "huhub." 




S.H. GROVE 



FIG. 9. Pinus caribaea. a, Leafy branch with attached cone ( X 1 A). b, Cone, 
unopened (X 1). 



47 



48 FIELDIANA: BOTANY, VOLUME 24 

This pine has been confused with the slash pine of the southeast- 
ern United States, P. Elliottii Engelm., which does not occur in Cen- 
tral America. From this species it may be distinguished by the 
shorter weaker prickles of the cone scales, the smaller seeds with 
wings usually remaining attached, and the grouping of leaves pre- 
dominantly in fascicles of 3's. In Journ. For. 50: 918-923. 1952, Drs. 
Little and Dorman discuss the taxonomy and nomenclature of the 
slash pine (P. Elliottii} and the Caribbean pine (P. caribaea). 

Pinus Montezumae Lambert, Descr. Pinus ed. 3. 1: 39. 1832. 
P. fiUfolia Lindl. Bot. Reg. 26, Misc. 61. 1839. P. Montezumae var. 
Lindleyi Loudon, Encycl. Trees 1004. /. 1882. 1883. Pino; Pino 
Colorado; Pino de ocote. Figure 10. 

Mountain slopes or plains, often forming extensive stands, at 
about 1050-3000 meters; Baja Verapaz; Chimaltenango; Quiche"; 
Huehuetenango; Totonicapan; Quezaltenango; San Marcos. Widely 
distributed in Mexico. 

A medium-sized or large tree, sometimes 30 meters high with a trunk 75 cm. 
or even more in diameter; bark dark gray, very rough, broken into conspicuous 
plates separated by deep furrows and ridges; youngest branches stout, 8-12 mm. 
thick, very rough, the decurrent bases of the sheaths of the leaf fascicles close to- 
gether and conspicuous; leaves usually in fascicles of 5, occasionally in 4's or 6's, 
thickly set on young branches, dark grass-green, the youngest leaves somewhat 
lighter and brighter green, 15-45 cm. long, 1-1.5 mm. thick, stout and usually 
rather stiff, erect, abundantly serrulate, with stomata on all sides, the resin ducts 
medial, 3 to 6, the leaf with 2 fibre-vascular bundles; leaf sheaths persistent and 
conspicuous; cones opening at maturity, deciduous or nearly deciduous, variable 
in size, ovoid-conic to long-conic, spreading or drooping, 18-30 cm. long or often 
shorter, 4.5-8 cm. thick, dull to rich brown; cone scales thickened at the tip; umbo 
dorsal, the prickle inconspicuous, mostly deciduous; wing of the seed thin, 2-2.5 cm. 
long, its base thin or slightly thickened. 

This is one of the common pines of western Guatemala, often 
growing in association with P. oocarpa and perhaps P. pseudostrobus. 
All these species are much alike in the field. 

Pinus Montezumae var. rudis (Endl.) Shaw, Pines Mex. 22. 
1909. P. rudis Endl. Syn. Conif. 151. 1847. P. Hartwegii Lindl. 
Bot. Reg. 25, Misc. 62. 1839. P. Montezumae var. Hartwegii Engelm. 
Trans. St. Louis Acad. Sci. 4: 177. 1880, according to many authors, 
but the combination not legitimately made. P. Donnell-Smithii 
Masters in Bonn. Smith, Bot. Gaz. 16: 199. pi. 2. 1891 (type col- 
lected near the summit of Volcan de Agua, Sacatepe"quez, J. D. 
Smith 2182). Teccamanil (San Marcos); Pino; Pino Colorado; Pino 
de ocote. 




FIG. 10. Pinus Montezumae. a, Leafy branch with attached cone ( X H) 
6, Fascicle of needles (X %). 



49 



50 FIELDIANA: BOTANY, VOLUME 24 

Occurring at high elevations, chiefly on the upper slopes of the 
volcanoes and on the high mountains of Huehuetenango, in a belt 
of about 3000-3500 meters elevation; Sacatepe"quez (Volcan de 
Agua); Chimaltenango (volcanoes of Fuego and Acatenango, Cerro 
Tecpan); Totonicapan; Huehuetenango (Sierra de los Cuchuma- 
tanes); Quezaltenango (Volcan de Santa Maria); San Marcos (vol- 
canoes of Tajumulco and Tacana). Widely distributed in Mexico. 

Usually a large tree, sometimes 30 meters high but usually lower, at the highest 
elevations often a shrub of 2 meters, the trunk usually tall and slender, the crown 
often dense and compact; leaves shorter than in the typical variety, mostly 7-15 
cm. long; cones generally shorter and darker, often dark purple or almost black, 
mostly 7-13 cm. long and 5-8 cm. thick. 

This is one of the most easily recognized of the pitch pines of 
Guatemala, because of its short leaves and the limited areas in which 
it grows. Although treated as a distinct species by some authors, 
such as Martinez and Loock, the differences by which it is supposed 
to be separated do not hold true in Guatemalan material. The leaves 
are nearly always erect in the Guatemalan trees of P. Montezumae. 
On some of the volcanoes it can be noted that trees of this species 
gradually become smaller and more dwarfed as the summit is ap- 
proached. With dwarfing of the trees, the leaves also become shorter. 
Thus, at the higher elevations, P. Montezumae var. rudis appears very 
distinct, but at lower elevations it is not easily distinguishable from 
typical P. Montezumae. The differences used by Loock (loc. cit. 
p. 94) to separate P. Hartwegii from P. rudis, based upon color of 
cone, type of apophyses, thickness of cone scale, and constant num- 
bers of leaves in a fascicle, certainly do not hold true in Guatemalan 
material examined. Both of these entities are therefore considered 
to be the same and are treated as P. Montezumae var. rudis. 

On the volcanoes as well as in the Cuchumatanes (where it grows 
on limestone), it forms forests of distinctive aspect, the trees of rather 
pale appearance as to foliage, and usually standing well apart in areas 
covered with dense stands of bunch-grasses. On the central and 
western volcanoes the variety marks a very distinct zone of vege- 
tation, beginning at the upper limit of the moist or wet, very dense, 
broad-leafed forest, where no pines occur, and extending to the sum- 
mits of the volcanoes. At highest elevations the plants often grow 
in the most exposed places, subjected to the strongest gales. The 
trees are progressively smaller as the elevation increases. Vast num- 
bers of dead trees occur on the upper slopes of the central volcanoes. 
The Indians insist they have been killed by gases from the craters, 
but it is probable that all of them are victims of insect attack. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 51 

Pinus oocarpa Schiede, Linnaea 12: 491. 1838. P. oocarpoides 
Lindl. ex Loudon, Encycl. Trees 1118. 1883. P. tecumumani Schwert- 
feger, Inf. Gob. Guat. sobre Entom. For. 1: 39. figs. 112-122, 139. 
1953 (without Latin diagnosis, type from Guatemala, Schwertfeger 
52). Pino; Pino de ocote; Chaj (Coban, Quecchi). Figure 11. 

The most abundant pine of Guatemala, widely distributed on 
mountain slopes and plains, sometimes forming almost pure stands 
but often associated with oaks and sometimes with other pines, occur- 
ring at 1000-2700 meters; Alta Verapaz; Baja Verapaz; El Progreso; 
Zacapa; Chiquimula; Jalapa; Jutiapa; Guatemala; Sacatepe"quez; 
Chimaltenango; Solola; Quiche"; Totonicapan; Huehuetenango; 
doubtless also in other departments. Widely distributed in Mexico; 
Honduras; El Salvador; Nicaragua. 

A large or medium-sized tree, 15-25 meters high or often taller, often with a 
thick trunk; bark grayish- or reddish-brown, rough, deeply fissured into broad 
plates; young shoots uninodal, pruinose; youngest branchlets 6-10 mm. thick, 
moderately or conspicuously rough, light or reddish-brown; leaves mostly in fasci- 
cles of 5, sometimes in 4's, rich olive-green or grass-green on all sides, 12-30 cm. 
long, 1-1.2 mm. thick, flexible and rather slender, subcoriaceous, ascending to erect, 
abundantly serrulate; stomata on all sides of the leaf, the resin ducts mostly septal, 
sometimes internal, the leaf with 2 fibro- vascular bundles; leaf sheaths persistent 
and conspicuous; cones broadly ovoid to ovoid-conic, persistent, opening late in 
maturity, the lower scales usually remaining closed for a long period, the cones 
pendulous, on a peduncle as much as 3.5 cm. long, light yellow-brown or reddish- 
brown; cone scales thickened at the tip, the umbo dorsal, often with a spine-like 
point; wing of the seed often thickened at the base. 

This is probably the most abundant of all the Guatemalan pines, 
being the commonest species of the central and northwestern moun- 
tains, where it forms very extensive stands in some regions. It is an 
important source of lumber. The species extends southward far into 
the interior of Nicaragua, which is the southern limit for the distri- 
bution of mountain pines in Central America. It is the common pine 
of the Coban region of Alta Verapaz, where the rainfall is much 
greater than in most parts of its range and the pine forests are excep- 
tionally thrifty and handsome. These forests, however, have been 
much depleted by lumbering operations. Here the pines usually 
grow in association with Liquidambar rather than oaks. The young 
plants about Coban often make an extraordinary growth, attaining 
a height of 2 meters before branching. The handsome, very pale, 
yellowish wood is much used about Coban for all kinds of construc- 
tion, and also for furniture. It supplies most of the lumber used in 
the region. 




S.H. Grove 



FIG. 11. Pinus oocarpa. a, Portion of leafy branch with attached cone ( X 
b, Cone, unopened (X 1). 



52 



I STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 53 

About 40 or 45 years ago Gustav Helmrich planted a large area 
on the hillsides of Finca Samac near Coban with the native pine. 
The trees now are medium-sized or large and ready to cut. From a 
distance they have every appearance of being a native forest. Oak 
trees planted at the same time are about half as large. 

The complexities of the taxonomy and nomenclature of Pinus 
species have been recently augmented by the publication of another 
species, Pinus tecumumani, proposed, without Latin diagnosis, by 
Dr. Fritz Schwertfeger (ibid. 1953). One of the specimens (Schwert- 
feger 52), upon which the type collection is based, was identified by 
the junior author as P. oocarpa Schiede. Despite Dr. Schwertfeger's 
insistence on the recognition of this and other collections which he 
made (nos. 38, 69, 70, 71, and 72) as constituting an undescribed 
species, a re-examination of his material does not support his claim. 
He himself admits (loc. cit., p. 39) that, from a botanical standpoint, 
P. tecumumani may be considered as a variety or form of P. oocarpa, 
and that the two resemble one another in many respects. 

Both P. oocarpa and P. pseudostrobus appear to be variable, and 
the type material of P. tecumumani seems best interpreted as repre- 
senting a somewhat variant form of P. oocarpa, although it is also 
possible that it may be considered as of hybrid ancestry between 
P. oocarpa and P. pseudostrobus. The shape, size, and short stipe of 
the cone would ally P. tecumumani to P. oocarpa, whereas the lesser 
thickness of the needles and their sometimes pendent position would 
ally P. tucumumani to P. pseudostrobus. 

Pinus pseudostrobus Lindl. Bot. Reg. 25, Misc. 63. 1839. P. 
tenuifolia Benth. PI. Hartweg. 92. 1842. P. pseudostrobus var. tenui- 
folia (Benth.) Shaw, Pines of Mexico 20. pi. 13. 1909. Pino; Pino de 
ocote; pino bianco and pinabete (Honduras.) Figure 12. 

Usually in rather moist forest, at 1000-3000 meters; descending 
on the Pacific slope to 850 meters, but in most parts of the country 
found at 1500 meters or higher; Alta Verapaz(?); Baja Verapaz; El 
Progreso; Jalapa; Guatemala; Sacatepe*quez; Chimaltenango; Solold; 
Suchitepe"quez; Quezaltenango; Totonicapan; Huehuetenango; per- 
haps in some other departments. Widely distributed in Mexico; 
British Honduras (Baker's Pine Ridge); Honduras; Nicaragua. 

A medium-sized or large tree, 10-30 meters high, the trunk sometimes almost 
a meter in diameter, the bark smooth and gray at first, becoming brownish-gray or 
blackish in age, the upper part of the trunk and the branches sometimes reddish- 
brown; youngest branches 4-8 mm. thick, almost smooth or slightly roughened, 
light to dark brown or reddish-brown, conspicuously pruinose, the internodes long; 



54 



FIELDIANA: BOTANY, VOLUME 24 




S.H. Grove 



FIG. 12. 
attached (X 



Pinus pseudostrobus. Branch, showing pendent leaves and cone 



leaves usually 5 in a cluster, sometimes 6, gray-green or grass-green on all sides, 
flexible, soft, slender, drooping or laxly spreading or arching, 12-35 cm. long, 0.6-1 
mm. thick, moderately and minutely serrulate; stomata present on all sides of the 
leaf, the resin ducts medial, the leaf with 2 fibro- vascular bundles; leaf sheaths 
persistent; cones dehiscent and deciduous at maturity, ovoid to ovoid-conic or 
oblong-conic, 5.5-14 cm. long, 2.5-10 cm. thick, chestnut to dark brown; cone 
scales thickened at the tip, sometimes more or less prolonged, the umbo dorsal, 
sometimes with a salient tip; wing of the seed articulate, its base thin or slightly 
thickened. 

This species is easily confused with P. oocarpa, from which it may 
be distinguished by the usually drooping, less rigid leaves, usually 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 55 

sessile cones which open at maturity, generally more slender, less 
roughened branches, and medial rather than septal resin ducts. It 
is a common pine of the central regions, forming sometimes pure 
stands but often mixed with other pines or with oaks. 

Although Martinez (loc. cit., p. 68) recognizes P. tenuifolia Benth. 
as a distinct species, we are unable to separate it from P. pseudostrobus, 
either specifically or varietally, as it was treated by Shaw. Martinez 
uses the character of the association of the cone with an oblique 
peduncle as the principal means of separating P. tenuifolia from P. 
pseudostrobus. We have not found this or other characters stressed 
by Martinez to hold true for use in distinguishing P. tenuifolia from 
P. pseudostrobus. 

Pinus Strobus L., var. chiapensis Martinez, Anal. Inst. Biol. 
Mexico 11: 81. 1940. Pinabete (Chiapas). 

Mountain slopes, in stands of pine and hardwood forests, gen- 
erally at elevations between 800-2000 meters; Quiche" (Rio Suchun 
below Nebaj) ; Huehuetenango (Sierra de los Cuchumatanes). South- 
ern Mexico (Vera Cruz, Oaxaca, and Chiapas). Figure 13. 

A large tree, sometimes 30 meters high with a trunk up to 1 meter or more in 
diameter; bark rough, light gray to brownish, broken into shallow furrows and 
ridges; younger branches smooth, grayish to greenish-gray; leaves 5 in a fascicle, 
light green or yellowish-green, silvery or whitish beneath, 5.5-12.5 cm. long, 0.6-1 
mm. thick, slender, flexible, serrulate, the teeth small and remote, the stomata 
ventral only, the resin ducts external, the fibro-vascular bundle only 1 ; leaf sheaths 
soon deciduous; cones broadly cylindrical, slightly tapering towards the apex, 
10-13 cm. long, 4.5-6 cm. thick, yellowish to reddish-brown, rather dull; cone scales 
flattened, the apophysis yellowish-brown, rounded and not prolonged at apex, the 
body reddish-brown, the umbo terminal; seeds dark brown, with a brown wing 
2.5 cm. long and 8-9 mm. wide. 

Like Pinus caribaea, this species is of very limited distribution in 
Guatemala, being confined to mountain forests in the departments 
of Quiche" and Huehuetenango. It resembles P. Ayacahuite, but 
occurs at lower elevations, generally between 1400 and 1800 meters, 
but sometimes at elevations as low as 800 meters, on the lower slopes 
of the Sierra de los Cuchumatanes. On Cerro Victoria, near Barillas, 
in the department of Huehuetenango, it is associated with forests of 
Liquidambar, while in the department of Quich its forest associates 
are Liquidambar, Acer Skutchii, Platanus, Quercus, Fraxinus, and 
others whose relationships are with the flora of the temperate zone of 
the southeastern United States. 



56 



FIELDIANA: BOTANY, VOLUME 24 




(x 



S.H.G 



FIG. 13. Pinus Strobus var. chiapensis. a, Leafy branch (X 



b, Cone 



TAXODIACEAE. Bald Cypress Family 



Reference: R. Pilger, Taxodiaceae, Pflanzenfamilien ed. 2. 13: 
342-360. 1926. 

Large trees; leaves scale-like or needle-like, sometimes falcate, persistent or 
deciduous; staminate flowers solitary, terminal or axillary, capitately clustered or 
in long pendent panicles; filaments short, the anther scale broad, with 2-9 free 
sporangia pendulous from its lower edge; pistillate inflorescence solitary, terminal, 
with numerous spirally arranged bracts, each bearing 2-9 ovules; scales of the cone 
ligneous or thick-coriaceous, rounded, opening at maturity; seeds with a narrowly 
winged margin. 

Nine genera, seven of them Asiatic. The only other American 
one is Sequoia, confined to the Pacific coast of the United States, 
whose species include the most massive trees known. 



: STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 57 

Leaves deciduous; staminate flowers in terminal, much elongate, drooping panicles. 

Taxodium 

Leaves persistent; staminate flowers in short stiff spikes at the ends of the branches. 

Leaves subulate, 6-8 mm. long Cryptomeria 

Leaves linear-lanceolate, mostly 25-40 mm. long Cunninghamia 



CRYPTOMERIA D. Don 

Evergreen trees with reddish-brown bark peeling off in long shreds, the crown 
pyramidal, with spreading branches; leaves spirally arranged, in 5 vertical ranks, 
decurrent at the base, directed forward and curved inward, awl-shaped, laterally 
compressed; flowers monoecious; staminate flowers oblong, composed of numerous 
imbricate stamens, sessile, axillary, forming short spikes at the ends of the branch- 
lets; pistillate flowers globose, solitary at the ends of the short branchlets; cones 
brown, subglobose, maturing the first year but persistent after shedding their seeds; 
cone scales 20-30, woody, cuneate, enlarged above into a disk, this bearing a 
recurved mucro at its middle and with 3-5 pointed processes on the upper margin; 
seeds 2-5 in each scale, triangular-oblong, slightly compressed, narrowly winged; 
cotyledons 2-3. 

The genus consists of a single species. 

Cryptomeria japonica (L. f.) D. Don, Trans. Linn. Soc. 18: 
167. 1841. Cupressus japonica L. f. Suppl. PI. 421. 1781. 

Native of China and Japan, occasionally planted for ornament 
about Guatemala City and probably elsewhere. 

A large tree, sometimes becoming 50 meters high, with a very thick trunk; 
leaves linear-subulate, rigid, subulate, 6-8 mm. long, laterally compressed, carinate 
above and beneath, with stomata on both sides, bright green; staminate flowers 
about 6 mm. long; cones globose, 1.5-2.5 cm. in diameter; seeds dark brown, 
5-6 mm. long. 

The tree is uncommon in Guatemala, but there are many fine 
large individuals in Costa Rica. It seems to thrive in mountain 
regions of the tropics, especially where there is abundant moisture. 



CUNNINGHAMIA R. Brown 

Evergreen trees, the branches spreading, pendulous at the ends; leaves spirally 
arranged, decurrent at the base, linear-lanceolate, 2-ranked, spreading; flowers 
monoecious, the staminate flowers cylindric-oblong, in terminal clusters; pistillate 
flowers globose, 1-3 at the end of a branchlet; cones subglobose, the scales thick- 
coriaceous, broadly ovate, pointed, irregularly serrate, each scale bearing 3 nar- 
rowly winged seeds; cotyledons 2. 

The genus consists of two species, both Asiatic. 



58 FIELDIANA: BOTANY, VOLUME 24 

Cunninghamia lanceolata Hook. Bot. Mag. pi. 2743. 1827. 

Native of southern, central, and western China; planted for orna- 
ment and shade in Guatemala City and probably elsewhere in the 
country. 

A large tree, attaining a height of 25 meters, the bark brownish, separating in 
irregular plates and exposing the reddish inner bark; leaves crowded, linear- 
lanceolate, spine-tipped, mostly 3-6 cm. long, lustrous above, with 2 white longi- 
tudinal bands beneath; cones globose-ovoid, 2.5-5 cm. long, brown. 

TAXODIUM L. Richard. Bald Cypress 

Resinous trees, usually deciduous, the bark light brown, shallowly furrowed 
and scaly, the branches suberect or spreading; branchlets of 2 kinds, those near the 
tips of the branches persistent and with axillary buds, those on the lower part of 
the branch deciduous and without axillary buds; winter buds globose, scaly; leaves 
alternate, subulate or flat and linear, with 2 bands of stomata on the lower surface, 
the leaves of the persistent branchlets spreading, deciduous; flowers small, mon- 
oecious; staminate flowers in slender drooping panicles, the flowers consisting of 
6-8 spirally arranged stamens; pistillate flowers solitary or geminate near the ends 
of the branches of the preceding year, subglobose, consisting of imbricate scales, 
each scale bearing 2 ovules within near the base; fruit a short-stalked, globose or 
ovoid, small cone, ripening the first year, consisting of many thick, spirally ar- 
ranged, peltate scales, these dilated from a slender stipe into an irregularly 4-sided, 
often mucronate disk, the fertile scales each with usually 2 seeds; seeds unequally 
trigonous, with 3 thick wings; cotyledons 4-9. 

One and perhaps two other species are known, in the southern 
and southeastern United States. 

Taxodium mucronatum Tenore, Ann. Sci. Nat. III. 19: 355. 
1853. T. mexicanum Carriere, Trait. Conif. 147. 1855. Sabino; 
Camphor eta, (Suchitepe"quez, where cultivated). 

Along the borders of streams, often growing in shallow water, 
800-2000 meters; Huehuetenango (extending eastward to Los Pini- 
tos, just southeast of Huehuetenango; collected also near Demo- 
cracia, near Santa Ana Huista, near Cuilco, and San Sebastian). 
Widely distributed in Mexico. Figure 14. 

A large tree, sometimes 30 meters high, the trunk tall, straight, enlarged near 
the base, the bark brownish-red or often very pale; leaves, and many of the branch- 
lets, deciduous, linear, 6-12 mm. long, thin and soft; cones subglobose, brown, 
1.5-2.5 cm. in diameter. 

In Mexico usually called "cipre"s." This is a common tree in parts 
of Mexico and there are some giant historic trees there, the most 
famous being that at Santa Maria del Tule, Oaxaca, which has a 
height of 38 meters and a trunk circumference of 51 meters, with a 




S.H. Grov* 

FIG. 14. Taxodium mucronatum. a, Flowering branch with leaves (X }/). 
b, Fruiting branch (X 1). c, Seed, dorsal view (X 3). d, Seed, ventral view 
(X 3). 

59 



60 FIELDIANA: BOTANY, VOLUME 24 

maximum diameter of 12 meters. The Cipre"s de Montezuma at 
Chapultepec has a height of 51 meters and a trunk circumference of 
15 meters. At the time of the Conquest it was a noted tree, and its 
age has been estimated at 700 years. No other trees of comparable 
size are found in Guatemala. At the southeastern limit of their 
range, near Huehuetenango, the trees are few and small, although it 
is probable that formerly they were more numerous and with larger 
individuals among them. Some trees grow along the stream at the 
base of the hill on which lie the ruins of Zaculeu, the site of the large 
Indian city that preceded the present city of Huehuetenango. It is 
probable that the name of this settlement is derived from the Nahuatl 
name for Taxodium, ahuehuetl, and at first probably was Ahuehuete- 
nango. A different origin of the name is given by Guatemalan 
writers, but the one here suggested seems fairly obvious. All the 
Indian cities had perfectly good names in the Indian tongues, but 
when the Spanish Conquistadores arrived, accompanied by their 
Mexican mercenaries, the latter renamed all the cities in their own 
language, and all or most of these names persist to the present time, 
in regions where Nahuatl dialects never were spoken by the original 
inhabitants. 

The wood of this bald cypress is light or dark brown or yellowish 
and susceptible of a good polish. It is soft and rather weak, but in 
Mexico it is esteemed for construction purposes, especially because 
it is resistant to decay and insect attacks. In Guatemala it is too 
rare to be of economic importance. 

The leaves of the bald cypress in Guatemala turn yellow and red 
about the first of January, or perhaps even earlier, and are then very 
conspicuous. The few scattered trees along the Rio Naranjo below 
Huehuetenango are poorly developed, but they were found fruiting 
abundantly, and there were many seedlings about them. Some of 
the trees grow in running water, which must be much deeper during 
the height of the rainy season. Scattered Taxodium trees are in cul- 
tivation in various parts of the country, as about the capital and 
Coban and even in the fincas between Colomba and Coatepeque. 

TAXACEAE. Yew Family 

Trees or shrubs with evergreen foliage, resinous in almost all parts; leaves alter- 
nate and usually 2-ranked or spirally arranged, linear, entire; flowers dioecious, the 
staminate flowers solitary or in small cones, axillary; stamens with 2-8 sporangia; 
pistillate flowers borne on small axillary twigs, the ovary terminal, surrounded at 
the base by pairs of scales; seeds partially enclosed in a fleshy colored aril; cotyle- 
dons 2. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 61 

Four genera are known, Taxus, Torreya, with 5 species in Japan 
and China, California, and Florida, Austrotaxus, with one species in 
New Caledonia, and Amentotaxus, with one species in China. 

TAXUS L. Yew 

Trees or shrubs, the bark reddish or reddish-brown, scaly, the branches usually 
spreading; branchlets irregularly alternate, the winter buds with imbricate scales; 
leaves spirally arranged, spreading in 2 ranks, linear, often falcate, with 2 broad, 
yellowish or grayish-green bands beneath, without resin ducts; flowers dioecious or 
rarely monoecious, the staminate in stipitate heads of 6-14 stamens, each stamen 
with 5-8 sporangia; pistillate flowers consisting of several imbricate scales, the 
terminal one bearing an ovule with a disk at its base; seeds ovoid, subangulate, 
surrounded by a red fleshy cup open at its apex; endosperm uniform. 

Seven or eight species are known, widely distributed in the north- 
ern hemisphere. They are all closely related, differing principally in 
range, and have been treated by some authors as forms of a single 
species, the European T. baccata L. Four species are known from 
North America, none in South America. The heartwood is bright 
orange to brown, the sapwood thin and white; of rather high luster, 
odorless and tasteless; hard, moderately heavy, stiff, and elastic, with 
a specific gravity of about 0.70; of very fine and uniform texture and 
straight or variable grain. The wood has long been used in Europe 
for bows, and at the present time its principal use is for the same 
purpose. 

Taxus globosa Schlecht. Linnaea 12: 496. 1838. Cipres; 
Pinabete. 

In cloud forest in the mountains at rather high elevations, 2200- 
3000 meters; Baja Verapaz; El Progreso (Sierra de las Minas; Volcan 
de Santa Luisa); Zacapa (Sierra de las Minas, Volcan Gemelos; 
Monte Virgen) ; Huehuetenango (Sierra de los Cuchumatanes, Cerro 
Canana). Southern Mexico (San Luis Potosi; Hidalgo; Veracruz; 
Oaxaca). Figure 15. 

A large shrub or a small or medium-sized tree, sometimes 15 meters high; leaves 
linear, somewhat falcate outwardly, 2-3.5 cm. long, 2-3 mm. wide, cuspidate, 
slightly narrowed at the sessile base, olive-green above and lustrous, somewhat 
silvery and pale beneath, the margins revolute in drying, the costa prominent be- 
neath and also on the upper surface; mature branchlets reddish-brown, somewhat 
angulate; pistillate branchlets rather stout; cup subtending the seed about 4 mm. 
broad (when dry); seed ovoid or oval-ovoid, about 5 mm. long, apiculate at the 
apex. 

This tree seems to be local in the Guatemalan mountains, al- 
though it is plentiful in the localities where it has been found. These 




FIG. 15. Taxus globosa. a, Leafy branch ( X Vi). b, Fruit (X 4). 

62 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 63 

are separated by a long distance from the nearest Mexican localities 
from which the tree has been reported. The seeds of the various 
yews are reported to contain a poisonous alkaloid (taxine) and eating 
them has sometimes caused the death of human beings. The bark is 
said to be rich in tannin. 



A species of Ephedra, of the family Ephedraceae, is in cultivation 
in the Jardin Botanico in Guatemala. It is a low shrub with naked, 
jointed branches, the leaves represented by small scales, the fruit 
small and cone-like. The determination of the plant is uncertain, 
but it may be the European E. distachya L. 

TYPHACEAE. Cat-tail Family 

References: P. Graebner, Typhaceae, Pflanzenreich IV. 8. 1900. 
Percy Wilson, Typhaceae, N. Amer. Fl. 17: 3-4. 1909. 

The family consists of a single genus. 

TYPHA. Cat-tail 

Glabrous herbs, growing in water or wet soil, with creeping rootstocks and sim- 
ple erect stems; leaves linear or strap-shaped, sheathing at the base, flat, more or 
less convex on the back, entire, parallel- veined ; flowers unisexual, densely crowded 
in compact cylindric spikes, the upper portion of the spike staminate, the lower 
pistillate; staminate inflorescence terminal, separated or contiguous to the pistil- 
late inflorescence; each spike usually subtended by a spathaceous fugacious bract 
in early anthesis, the flowers irregularly intermingled with variously shaped hairs; 
stamens 1-7; filaments short or long, free or connate; anthers linear or oblong, 
basifixed, 2-celled, longitudinally dehiscent, the connective produced beyond the 
cells in a conic fleshy acumen; pollen grains simple or compound; pistillate flowers 
ebracteolate or mingled with slender, clavate or spatulate bractlets, and often with 
abortive pistillate flowers mixed with them ; gynophore slender, the perianth com- 
posed of several delicate, silky, simple or clavate hairs; ovary superior, usually 
1-celled, more or less stalked, fusiform, with a solitary pendulous anatropous ovule; 
style elongate, slender, erect; stigma linear, spatulate, or rhombic-fusiform, uni- 
lateral; fruit minute, subsessile or long-stipitate, fusiform or ellipsoid, with a 
membranous or coriaceous pericarp, longitudinally splitting; seed subcylindric or 
narrowly ellipsoid, the testa thin, membranaceous; endosperm carnose or farina- 
ceous; embryo cylindric, straight. 

About nine species, of wide distribution. Only the following are 
known from Central America. 

Staminate and pistillate portions of the spike contiguous; pistillate spike dark 
chocolate-brown; pistillate flowers ebracteolate; stigma flattened, obliquely 
rhombic-lanceolate; aborted pistil clavate; pollen grains in 4's, 4-celled. 

T. latifolia 



64 FIELDIANA: BOTANY, VOLUME 24 

Staminate and pistillate portions of the spike separated; pistillate spikes light or 
reddish brown; pistillate flowers bracteolate; stigma linear or narrowly lan- 
ceolate; pollen grains single, one-celled T. domingensis 

Typha latifolia L. Sp. PI. 971. 1753. Tul 

In shallow water, especially at the edges of lakes, 1000-1900 
meters; Alta Verapaz; Baja Verapaz; Sacatepe*quez (near Antigua); 
Solola (Finca Moca) ; Huehuetenango. Widely distributed in North 
America and in the Old World. 

Plants robust, 1-2.5 meters tall; leaves broadly linear and surpassing the in- 
florescence, flat, 6-25 mm. wide; staminate and pistillate spikes about equal in 
length; staminate inflorescence mustard or brownish-yellow, 7-13 cm. long, decid- 
uous after flowering, the hairs sordid white, linear, acute at the apex; pistillate 
inflorescence 0.5-3.5 cm. thick, 7-30 cm. long, at least 6-8 (up to 20) times longer 
than broad; stigma dark brown or blackish, surpassing the hairs of the gynophore; 
denuded rachis of the mature pistillate spike stout, 8-11 mm. thick, conspicuous 
by the long bristle-like persistent pedicels. 

Recent studies reveal that the fiber of the cat-tails may be used 
as a substitute for cotton and linen in making articles of clothing, 
rugs, and many other products. The mature fruiting spikes are often 
used in the United States for their ornamental value, and the solid 
and young spikes are pickled and served in the form of a relish or 
in salads. 

Typha domingensis Pers. Syn. PI. 2: 532. 1807. T. truxillensis 
HBK. Nov. Gen. & Sp. 1: 82. 1815. T. angustifolia of many authors, 
not L. Tul; Tule; Espadana; Enea. Figure 16. 

In wet soil or often in shallow water, in ditches or along the bor- 
ders of lakes, ponds, and swamps, 1200 meters or lower; Izabal; 
Zacapa; Jutiapa; Guatemala; Quiche*. Widely distributed in the 
western hemisphere. 

Plants robust, 2-4 meters high; leaves flat, pale green, coriaceous, 5-20 cm. 
wide, usually not equaling the inflorescence; axis of the staminate inflorescence 
provided with reddish-brown hairs, these mostly branched, dilated at the apex, the 
branches curved; staminate inflorescence 0.7-2 dm. long; pistillate spikes pale 
brown, 10-40 cm. long, 5-22 mm. thick, increasing in thickness with approaching 
maturity; pedicels of the pistillate flowers up to 1 mm. long, the bractlets rhom- 
boid-, obcordate-, obovate-, or elliptic-spatulate, about as long as the hairs; hairs 
simple, slightly enlarged or thickened, spatulate at the brownish apex, shorter than 
the stigmas; denuded rachis of the mature pistillate spike slender, 3-4.5 mm. thick, 
merely roughened by the short rigid pedicels. 

The Maya name of Yucatan is "puh." This species has usually 
been confused with T. angustifolia, which is not known to occur in 




FIG. 16. Typha domingensis. a, Pistillate flower (X 8). 6, Inflorescence 
(X K). c, Staminate flower (X 10). d, Leaf (X >i). 



65 



66 FIELDIANA: BOTANY, VOLUME 24 

Mexico or South or Central America. True T. angustifolia may be 
distinguished by the shorter staminate spikes, the much darker, cas- 
taneous or reddish brown, usually more slender pistillate spikes, 
which are overtopped by the fewer darker green, more membrana- 
ceous, plano-convex leaves, and the lack of conspicuous bractlets on 
the surface of the pistillate inflorescence; T. domingensis has lighter 
brown pistillate spikes becoming thicker in age and surpassing in 
height the more numerous, paler green, coriaceous, flattened leaves, 
and the surface of the pistillate spikes is covered by the ovate blades 
of the bractlets intermixed with the stigmas. 

Some authors have interpreted Persoori's original publication of 
the name T. domingensis as intended by him to represent a cate- 
gory below that of species. Graebner, for example, in his treatment 
of the Typhaceae in Das Pflanzenreich (IV. 8. 14) interprets Per- 
soon's name as a subspecies of T. latifolia, although Graebner uses 
the name T. domingensis in specific rank. Urban, however, in his 
Symbolae Antillanae (8: 5. 1920) ascribes the first specific combina- 
tion of the name T. domingensis to Kunth (Enum. 3: 92. 1841), 
interpreting Persoon's name to be published as a subspecies. If Per- 
soon had actually published T. domingensis as a variety or subspecies, 
as might at first glance be supposed, since the name and description 
were inserted between his species no. 1 (T. latifolia) and species no. 2 
(T. media) and marked by an asterisk, then, under the present rules 
of nomenclature, the name would have to be rejected as a species in 
favor of the next specific name, i.e. T. truxillensis HBK., published 
in 1815. However, in the preface of volume 1, Persoon states that 
obscure or doubtful species are marked by a cross sign or asterisk 
("Speciebus obscuris, aut quoad sedem dubiis, vel accuratiori in- 
dagationi subjiciendis, signa crucis seu asteriscum apposui"). There- 
fore, the name T. domingensis, marked by an asterisk and thought 
by Persoon to represent a doubtful entity, was published as a bino- 
mial in specific rank, and as the first published binomial in that rank 
must be accepted as a validly published specific name over the later 
published T. truxillensis. 

In western North America the thick rootstocks were formerly 
used as food by some of the Indians. In Guatemala as well as in 
some other parts of Central America the fluffy "wool" from the 
flower spikes is used for stuffing pillows and cushions. It is not very 
satisfactory for the purpose, since it mats into hard lumps that are 
most uncomfortable in the case of pillows. The spongy leaves are 
much used for weaving the mats called petates tules, and for making 
sopladores, the fans used to fan charcoal fires. The plant is especially 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 67 

abundant about Lago de Amatitlan, where there are wide and dense 
stands, of considerable economic importance locally. The cat-tail is 
plentiful also in the north coast. There, especially in thin forest, the 
plants are fully 2 meters high. In the Laguna de Ocubila near 
Huehuetenango a curious phenomenon was observed. Most of the 
plants were about a meter high and all of approximately the same 
height. In the center of these large colonies were smaller ones of 
plants just twice as high and with slightly broader leaves. No inter- 
mediates were to be seen. It is possible that both T. domingensis 
and T. latifolia were represented, but it appeared that all the plants 
were of the same species. 

Typha glauca Godr. (Fl. Loire ed. 2: 20. 1843) has been reported 
from Guatemala. It is supposed to differ, like T. angustifolia, from 
T. domingensis in having fewer than 10 leaves, which are green and 
convex on the back and exceed the reddish-brown flowering spikes, 
in having shorter staminate inflorescences, and in the inconspicuous 
bractlets of the pistillate inflorescence. It is somewhat intermediate 
in characters between T. domingensis and T. angustifolia on the one 
hand, and between T. latifolia and T. angustifolia or T. domingensis 
on the other, and has been considered by many European workers to 
be of hybrid origin and was treated by Graebner in Das Pflan- 
zenreich (IV. 8. 10: 16. 1900) as a hybrid between T. angustifolia and 
T. latifolia. Dr. Norman C. Fassett likewise is of the opinion that 
T. glauca can be considered only as of hybrid origin between T. lati- 
folia and T. domingensis. 

PANDANACEAE. Screw-pine Family 

Reference: O. Warburg, Pandanaceae, Pflanzenreich IV. 9: 1-97. 
1900. 

Trees or shrubs, often scandent, the main stem often branched, frequently 
emitting roots; leaves mostly lanceolate or linear, sessile and sheathing at the base, 
spirally arranged, longitudinally nerved and plicate, usually spiny-margined; flow- 
ers dioecious, the inflorescence simply spicate or subcapitate or compound, usually 
subtended by colored spathe-like bracts, the flowers sessile and commonly densely 
crowded, without a perianth; stamens few to many, spicately or umbellately dis- 
posed, the filaments usually filiform, short or elongate, the anthers 2-celled, ovate 
to linear, dehiscent by longitudinal slits, sometimes apiculate, rarely appendaged; 
ovary of 1-many carpels, 1-many-celled, the carpels irregularly disposed or annular 
or biseriate; ovules solitary and attached laterally, or numerous on lateral pla- 
centae, ascending, anatropous, borne on long or short funicles; styles usually none, 
sometimes spiniform, conic, furcate, or dentiform; stigmas appressed to the apex 
of the carpel or sometimes erect, usually reniform or hippocrepiform, rarely linear 



68 FIELDIANA: BOTANY, VOLUME 24 

or oblong; fruit drupaceous or baccate, 1-many-celled; seed with a thin or crusta- 
ceous testa, the endosperm copious, oily, the embryo minute, basal. 

An Old World family of three genera. 

PANDANUS L. Screw-pine 

Trees or shrubs, usually erect, the trunk often covered with the sheaths of old 
leaves, branched or rarely simple; leaves linear, usually spiny-margined and long- 
sheathing; staminate spadices spicate and compound, the pistillate terminal, spi- 
cate or racemose, the bracts usually whitish; stamens numerous, the filaments free, 
the anthers linear or oblong, basifixed, erect; carpels 1-many; stigmas usually reni- 
form or hippo crepiform; ovules solitary; fruit a syncarp, globose, ovoid, or ellipsoid, 
rarely cylindric, the drupes densely crowded, the pericarp thin, the apical portion 
sometimes separating as a cap, the mesocarp fibrous or fibrous-fleshy; endocarp 
osseous, 1-many-celled; seed erect, commonly ovoid or fusiform. 

About 150 species in the Old World tropics, some of them introduced into culti- 
vation in the American tropics as ornamental plants. 

Pandanus tectorius Soland. ex Parkinson, Journ. Voy. En- 
deavour. 1773. P. odoratissimus L. f. Suppl. PL 424. 1781. P. odo- 
ratus Salisb. Prodr. 3. 1796. 

Planted abundantly in hedges and for ornament in the Pacific 
boca costa, and also for ornament generally in the lowlands, and 
even as high as Guatemala (1500 meters). Polynesia, southern Asia, 
and northern Australia. 

Plants shrubby or tree-like, sometimes 6 meters tall but in cultivation usually 
lower, the trunk emitting aerial roots; leaves greatly elongate, long-attenuate at 
the apex, the margins densely armed with slender sharp spines; staminate inflores- 
cence compound-spicate, very fragrant; stamens racemose-fasciculate; fruit as large 
as a human head, pendulous, the drupes 50-80 in the syncarp, red at maturity, 
4-10 cm. long. 

Occasional plants may be found in many of the lowland parks and 
fincas, but in some of the coffee regions, particularly in Suchite- 
pe"quez about Chicacao, and in Retalhuleu and Quezaltenango, along 
the road between Asintal and Colomba, there are long stretches of 
dense hedges of Pandanus. The form planted here has leaves with 
cream-colored margins. Perhaps also in cultivation is P. dubius 
Spreng., which has leaves 12-16 cm. wide (they are mostly 5-6 cm. 
in P. tectorius), abruptly contracted and cuspidate at the apex. 

POTAMOGETONACEAE. Pondweed Family 

Reference: Norman Taylor, Zannichelliaceae, N. Amer. Fl. 17: 13- 
27. 1909; Cymodoceaceae, loc. cit. 31-32. 



I STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 69 

Plants aquatic, glabrous, growing in fresh or salt water, the stems usually very 
slender, simple or branched; leaves petiolate or sessile, capillary or expanded into 
a distinct, narrow or broad blade, entire; flowers very small, perfect or monoecious, 
in sessile or pedunculate spikes, or clustered in the leaf axils; perianth none, the 
flowers sometimes enclosed in a hyaline sheath; androecium of 1-4 stamens; anthers 
extrorse, 1-2-celled, the connective sometimes petaloid; gynoecium of 1-4 distinct 
1-seeded carpels; fruits usually nut-like or drupe-like, sessile or stipitate; endo- 
sperm none. 

Genera about 9, the species widely distributed in tropical and 
temperate regions. Except for Potamogeton, the genera are all small, 
with only 1-6 species each. At least one other member of the family, 
Zostera marina L., has been found in Central America, in salt water 
along the coast of Honduras, and it is to be expected along the coasts 
of Guatemala. Zannichellia also might well be found in Guate- 
mala. 

Gynoecium of one carpel Diplanthera 

Gynoecium of several distinct carpels. 
Flowers umbellate, on long slender pedicels; stamens 2; connective of the anther 

not dilated; leaves narrowly linear Ruppia 

Flowers spicate, sessile; stamens 4; connective of the anther dilated and perianth- 
like; leaves almost filiform to oblong or oval Potamogeton 



DIPLANTHERA Thouars 

Submerged marine perennial plants, forming colonies with creeping rootstocks, 
rooting at the nodes of the articulated rootstocks; leaves linear, grass-like, more or 
less sheathing at the base; flowers dioecious; perianth none; staminate flower con- 
sisting of 2 anthers attached to the end of a long pedicel, the pedicels sheathed at 
the base; anthers unequally attached, one higher than the other, 2-celled; pistillate 
flowers consisting of a single naked carpel; style short, crowned with a slender, 
solitary stigma; mature fruit small, globose. 

Only one species is known. 

Diplanthera Wrightii (Aschers.) Aschers. in Engler & Prantl 
Nat. Pfl. Nacht. 37. 1897. Halodule Wrightii Aschers. Sitz.-ber. 
Ges. Nat. Freunde Berlin 1868: 19. 1868; Bot. Zeit. 26: 511. 1868. 

Submerged in salt water adjacent to the seashore, at sea level; 
Izabal (Bay of Santo Tomas and vicinity of Punta Palma near Puerto 
Barrios) . Southeastern United States (North Carolina to Florida) ; 
Mexico; West Indies. 

Leaves linear, flat, acute, ligulate at base, 3-40 cm. long, less than 1 mm. wide; 
stipular sheaths scarious, scarcely 3 cm. long; staminate flower of 2 anthers at the 
end of a long pedicel; anthers 2-celled, about 6 mm. long; pistillate flower of 1 naked 
carpel about 3 mm. long; style short; stigma solitary, slender; mature fruit black. 



70 FIELDIANA: BOTANY, VOLUME 24 

POTAMOGETON L. Pondweed 

References: J. 0. Hagstrom, Critical researches on the Potamo- 
getons, Svensk. Vet. Akad. Handl. 55, no. 5: 1-281. /. 1-118. 1916. 
M. L. Fernald, The linear-leaved North American species of Potamo- 
geton, section Axillares, Mem. Amer. Acad. 17: 1-183. pis. 1-40. 1932. 
E.G. Ogden, The broad-leaved species of Potamogeton of North Amer- 
ica north of Mexico, Rhodora 45: 57-105, 119-163, 171-216. illus. 
1943. 

Plants perennial, submerged or floating, the stems simple or branched, the ends 
of the stems often producing propagating buds; leaves usually of two kinds, float- 
ing and submerged, narrow or broad; floating leaves usually rather thick, generally 
expanded into a true blade; submerged leaves sometimes reduced to terete phyl- 
lodia but usually with broad or very narrow blades; stipules adnate to the leaf 
bases or free from them; inflorescences a simple or rarely branched spike, the 
flowers very small, green, crowded or the spikes sometimes interrupted; perianth 
none; stamens 4; anthers 2-celled, the connective enlarged and petaloid; carpels 4, 
distinct, sessile; ovules solitary in each cell; fruit an indehiscent nutlet; embryo 
completely or partially coiled into a circle. 

About 60 species, widely distributed, chiefly in temperate regions, 
few in tropical areas. Only the following species are known definitely 
from Central America. 

Leaves broad, all or most of them 1-1.5 cm. wide. 

Leaf blades cordate and clasping at the base P. perfoliatus 

Leaf blades acute to long-attenuate at the base P. illinoensis 

Leaves very narrow, all or most of them 3 mm. wide or narrower. 

Stipules adnate to the leaf bases P. peclinatus 

Stipules free from the leaf bases. 

Leaves usually without basal glands; peduncles usually 1 cm. long or shorter; 
fruiting spikes subcapitate or densely short-cylindric, 2-6-flowered; fruits 

with a conspicuous acute dorsal keel P. foliosus 

Leaves often with a pair of basal glands; peduncles mostly 1.5 cm. long or 
longer; fruiting spikes interruptedly cylindric, with 2-5 remote whorls of 
flowers; fruits rounded dorsally or very obscurely keeled P. pusillus 

Potamogeton foliosus Raf. Med. Rep. Hex. 2, 5: 354. 1808. 

At 900 meters; Solola (Finca Moca, J. Bequaert 53). Widely dis- 
tributed in North America; Mexico; West Indies. 

Plants wholly immersed, the stems very slender, a meter long or less, freely 
branched; leaves narrowly linear, slightly attenuate to the base, acute or subacute, 
usually without basal glands, the primary ones 0.3-2.7 mm. wide, 1-5-nerved; 
stipules forming tubular, delicately fibrous, blunt sheaths, 3-18 mm. long; pedun- 
cles slightly thickened above, mostly 3-10 mm. long; spikes subcapitate or thick- 
cylindric, at maturity 2-5 mm. in diameter, usually with 2-3 approximate whorls 
of 2 flowers each; fruits obliquely suborbicular or obovoid, compressed, 1.8-2.5 mm. 
long, with a thin or acute, undulate or strongly dentate dorsal keel. 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 71 

The single known Guatemalan collection was determined and 
cited by Fernald in the monograph cited above. 

Potamogeton illinoensis Morong, Bot. Gaz. 5: 50. 1880. P. 
fragillimus Hagstr. Svensk. Vet. Akad. Handl. 55, no. 5: 202. 1916 
(type from Laguna de Ayarza, Santa Rosa, Heyde & Lux). Yac- 
chian (Huehuetenango). 

Submerged or floating in slow streams or in lakes or ponds, 2400 
meters or lower; Pete"n; Alta Verapaz; Izabal; Jalapa; Jutiapa; Santa 
Rosa; Guatemala; Solola; Huehuetenango. Widely distributed in 
temperate North America; Mexico; West Indies. 

Stems slender, simple or branched; submersed leaves thin, elliptic or oblong- 
elliptic to lanceolate or linear, the blades 5-20 cm. long, 1.5-4 cm. wide, sessile or 
petiolate, acute and usually somewhat mucronate at the apex, 9-17-nerved; float- 
ing leaves present or absent, somewhat coriaceous, elliptic or oblong-elliptic, 4-13 
cm. long, 2-6.5 cm. wide, obtuse and mucronate at the apex, cuneate or rounded 
at the base, on petioles 2-9 cm. long, 13-29-nerved; stipules persistent, divergent 
and conspicuous, obtuse, those of the submersed leaves 2.5-8 cm. long; peduncles 
4-15 cm. long; spikes at anthesis of 8-15 whorls of flowers, in fruit cylindric and 
crowded, mostly 3-6 cm. long, 8-10 mm. thick; fruits obovate to orbicular or ovate, 
2.7-3.5 mm. long, conspicuously carinate. 

The plant has been reported from Guatemala as P. lucens L., 
P. Zizii Roth, and P. malaianus Miq. It is infrequent but some- 
times locally abundant. Apparently it varies from season to season 
in a given locality. In 1939 the senior author found it abundant in 
a small sluggish black stream near San Cristobal, Verapaz, at a time 
when it was impossible to reach the plants with any means at hand. 
In 1941, at the same season, he returned to the spot intending to 
procure specimens, but was unable to find a single plant where two 
years before the species had been so plentiful. 

Potamogeton pectinatus L. Sp. PI. 127. 1753. 

In water of lakes, 1200-1800 meters, or lower. Guatemala (Lago 
de Amatitldn) ; Solola (Lago de Atitldn). Widely distributed in tem- 
perate regions of both hemispheres; Mexico; British Honduras. 

Stems slender, submerged, usually much branched; leaves all submerged, linear 
or capillary, usually very numerous and often fasciculate, mostly 2-15 cm. long and 
3 mm. wide or less, rounded or obtuse at the apex and mucronate, 1-5-nerved; 
stipules adnate to the leaf bases, pale, the sheath 1-2 cm. long, the free portion of 
the stipules shorter, scarious; spikes long-pedunculate, the peduncles mostly 5-12 
cm. Jong; spikes 1-4 cm. long, usually much interrupted, with numerous whorls of 
flowers, the whorls sometimes 1 cm. apart; fruits somewhat 3-carinate, 2.5 mm. 
long or even larger. 



72 FIELDIANA: BOTANY, VOLUME 24 

Two forms of this species are found in Guatemala. That of Lago 
de Atitlan is the ordinary widespread form of the species. The plant 
of Amatitlan has much broader leaves, giving the plant a decidedly 
different appearance, and some of these specimens have been deter- 
mined by Fernald as P. striatus Ruiz & Pavon. Hagstrom considers 
this referable to P. pectinatus, calling it var. striatus (Ruiz & Pavon) 
Hagstrom of that species. As represented in Guatemala the two 
plants are so unlike that it seems probable that Fernald is correct in 
treating P. striatus as a distinct species. 

Potamogeton perfoliatus L. Sp. PI. 126. 1753. P. bupleuroides 
Fernald, Rhodora 10: 46. 1908. P. perfoliatus var. bupleuroides Far- 
well, Amer. Midi. Nat. 8: 264. 1923. 

At about 1800 meters; Solola (Lago de Atitlan). Widely distrib- 
uted in temperate North America; Eurasia. 

Stems elongate, often much branched, submerged; leaves all submersed, thin 
and delicate, sessile, orbicular to ovate-lanceolate, mostly ovate, 1-6 cm. long, 
0.5-3 cm. wide, 7-21-nerved, cordate and clasping at the base, rounded or very 
obtuse at the apex; stipules delicate, fugacious, ovate-oblong, rounded at the apex, 
2 cm. long or shorter; peduncles 1-9 cm. long, about as thick as the stems; spikes 
with 2-8 whorls of flowers, not crowded at anthesis, in fruit 1-2 cm. long and 8 mm. 
thick; fruits obovate, rounded dorsally, plump when mature, about 2.5 mm. long, 
with a prominent beak 0.7 mm. long. 

The Guatemalan material is referable to var. bupleuroides. 

Potamogeton pusillus L. Sp. PL 1: 127. 1753. Potamogeton 
panormitanus Bivona-Bernardi, Nuov. Piant. 6. 1938. P. panormi- 
tanus var. major G. Fischer, Bericht. Bayer. Bot. Gesell. 11: 109. 
1907. 

In ponds or small lakes, 2400 meters or lower; Quiche"; Huehuete- 
nango. Widely distributed in temperate regions of both hemispheres; 
southern Mexico; British Honduras; Cuba. 

Stems capillary, sometimes a meter long or more, usually much branched, the 
branches and their short lateral branchlets often terminated by winter buds; leaves 
linear or linear-setaceous, usually with a small translucent gland at the base, 1-7 
cm. long, 3 mm. wide or narrower, acute or obtuse, generally 3-nerved; stipules 
scarious-membranaceous, slender-tubular, their margins united to above the mid- 
dle, 6-17 mm. long; peduncles filiform throughout, 1.5-8 cm. long; spikes elongate, 
much interrupted, 6-12 mm. long, of 3-5 distant few-flowered whorls; fruits light 
olivaceous, obliquely obovoid, usually somewhat sigmoid, 2-2.8 mm. long, often 
deeply impressed on the somewhat flattened sides, the dorsal surface with a very 
low and broad, obscure keel. 



f STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 73 

RUPPIA L. 

Plants perennial, submerged, growing in salt or fresh water, the stems very 
slender, simple or usually branched; leaves alternate, linear, entire, 1-nerved, acute, 
slightly dilated and sheathing at the base; stipular sheaths clasping the leaf bases; 
flowers perfect, terminating a spadix-like peduncle; perianth none, the flowers often 
enclosed in the sheathing leaf bases at first; stamens 2, the anthers sessile, 2-celled; 
carpels of the ovary 4, sessile at first, becoming stipitate; stigmas sessile or at the 
end of a long or short style, peltate; mature fruits terminating a long slender pedi- 
cel, the pedicels umbellate, recoiling after the fruit falls; fruits oblique or sym- 
metric, crowned by the style; embryo curved, ovoid. 

Species 1, or perhaps more numerous, widely dispersed in tem- 
perate and tropical regions. 

Ruppia maritima L. Sp. PI. 127. 1753. 

Forming large submerged beds in shallow, salt or brackish water, 
at sea level, or in fresh-water lakes at an elevation of 1800 meters. 
Izabal (Rio Dulce; Bahia de Santo Tomas) ; Solola (Lago de Atitlan) ; 
Retalhuleu (Champerico). Mexico; British Honduras; El Salvador; 
Honduras; Costa Rica; widely distributed in temperate and tropical 
regions of both hemispheres. 

Plants often densely branched; leaves narrowly linear or almost filiform, 2-10 
cm. long, scarcely 0.5 mm. wide; stipular sheaths 6-10 mm. long; flowers on a short 
pedicel at first, this elongating in age and forming a loose spiral; stamens without a 
filament, early deciduous; mature carpels ovoid, often gibbous and somewhat 
oblique, 2 mm. long; style short and stout or somewhat attenuate, straight or unci- 
nate; pedicels of the fruits becoming 1.5-3 cm. long. 

One Guatemalan collection reported as Zannichellia palustris L. 
is actually referable rather to Ruppia maritima. 



NAJADACEAE 

Reference: Norman Taylor, N. Amer. Fl. 17: 33-35. 1909. 

Submerged aquatic herbs of fresh or salt water, with fibrous roots and very 
slender, branched stems, the internodes sometimes spinulose; leaves alternate, 
opposite, or verticillate, linear, spinose-dentate, sometimes mucronate, sheathing 
at the base; sheaths rounded or auriculate; flowers minute, monoecious or dioe- 
cious, solitary in the leaf axils; staminate flowers with a double perianth-like enve- 
lope, the inner hyaline, the outer 4-horned or entire; stamen 1, sessile, the anthers 
1-4-celled; pistillate flowers consisting of a single pistil, the style short or wanting, 
the stigmas 2-4, sometimes mingled with 1-3 sterile subulate stigmoid processes; 
fruit a sessile drupelet, its solitary seed filling the pericarp; embryo straight. 

The family consists of a single genus. 



74 FIELDIANA: BOTANY, VOLUME 24 

NAJAS L. 

A small group of about 10 species, generally distributed in tem- 
perate and tropical regions of both hemispheres. One other species, 
N. marina L., with relatively broad and deeply dentate leaves and 
dioecious flowers, has been collected in El Salvador, and is to be 
expected in Guatemala. Among Central American aquatic plants 
with linear leaves, the local species may be recognized at once by 
their toothed leaf margins. 

Leaves almost entire or very finely serrulate with minute subulate teeth. 

N. guadalupensis 
Leaves conspicuously serrulate with triangular teeth N. Wrightiana 

Najas guadalupensis (Spreng.) Morong, Mem. Torrey Club 3, 
pt. 2: 60. 1893. Caulinia guadalupensis Spreng. Syst. Veg. 1: 20. 
1825. 

In lakes, ponds, or small pools, sometimes in running water, 
ascending from near sea level to about 2400 meters; Pete"n; Zacapa; 
Chiquimula; Jalapa; Jutiapa; Escuintla; Guatemala; Solola; Quiche"; 
Huehuetenango. United States and Mexico to British Honduras 
and El Salvador; West Indies. 

Stems slender and much branched, often forming dense tangled masses; leaves 
all submerged, 10-25 mm. long, 0.5-1.5 mm. wide, acute or obtuse, usually tipped 
with 1-2 small spines, sheathing at the base but not auriculate, the marginal teeth 
small and inconspicuous, spinulose; flowers monoecious, the staminate 2-3 mm. 
long, with 4-celled anthers; pistillate flowers 2-3 mm. long; mature fruit tipped 
with 2-3 stigmas and usually with 1-2 spinulose sterile stigmoid processes, thus 
appearing to have 3-5 stigmas; seed ellipsoid, reticulate, with numerous quadran- 
gular areolae. 

Najas Wrightiana A. Br. Sitzb. Ges. Naturf. Fr. Berl. 17. 1868. 
N. multidentata Koch, Ber. Schweiz. Bot. Gesell. 44: 341. 1935 (type 
from Punta Gorda, British Honduras, W. A. Schipp 991}. N. 
Wrightiana subsp. Wrightiana R. T. Clausen, Bull. Torr. Bot. Club 
73: 364. /. d. 1946. N. Wrightiana subsp. multidentata R. T. 
Clausen, loc. cit. 365. 

In lakes, swamps, small ponds, and sometimes in running water, 
at 300 meters or less; Pete'n; Huehuetenango (Cie'naga de Lagartero). 
Eastern Mexico; British Honduras; West Indies; Brazil. 

Stems slender and much branched, often forming dense tangled masses; leaves 
all submerged, in dense fascicles, widely spreading, 10-25 mm. long, 0.4-0.9 mm. 
wide, acuminate, tipped with a single spine, conspicuously sheathed with a broad 
rounded base, the marginal teeth conspicuous, triangular, 15-40; flowers monoe- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 75 

cious, the staminate with a single 4-celled anther; pistillate flowers ovoid; mature 
fruit tipped with 2 stigmas and without sterile stigmoid processes; seed narrowly 
ellipsoidal, reticulate with numerous quadrangular areolae, 1-1.4 mm. long. 

According to Clausen's treatment, all the material from British 
Honduras and Guatemala, with 25-40 teeth on the margin of the 
relatively broader leaves, is included under Najas Wrightiana subsp. 
multidentata, while the plants of Cuba and the Mexican coastal plain 
fall under Najas Wrightiana subsp. Wrightiana. 

ALISMATACEAE. Water-plantain Family 

References: Fr. Buchenau, Alismataceae, Pflanzenreich IV. 15: 1- 
66. 1903; J. K. Small, Alismaceae, N. Amer. Fl. 17: 43-62. 1909. 

Herbs, aquatic or growing in wet soil, mostly perennial, lactescent; leaves 
usually all basal, petiolate, sheathing at the base, the blades linear to oval, hastate 
or sagittate, parallel-nerved; flowers perfect or polygamous, very rarely dioecious, 
borne upon a scape, usually pedunculate and arranged in bracteate whorls; peri- 
anth 6-parted, usually with a distinct division into calyx and corolla; sepals 3, free, 
imbricate in bud, generally persistent; petals 3, imbricate in bud, thin and delicate, 
white, fugacious; stamens 3 or multiples of 3, verticillate or spirally inserted, the 
filaments filiform or dilated; anthers 2-celled, affixed at or near the base, dehiscent 
by lateral longitudinal slits or sometimes extrorse; carpels 3 or in multiples of 3, 
verticillate or capitate upon a convex receptacle, free, 1-celled, usually 1-ovulate; 
styles terminal, the stigmas usually capitate; fruits as many as the ovaries, free, 
mostly 1-seeded, coriaceous, indehiscent; seeds mostly solitary and basilar, ascend- 
ing, with thin testa, smooth or rugose; endosperm none, the embryo hippocrepiform. 

Twelve genera are known, widely dispersed in temperate and 
tropical regions of both hemispheres. Only the following genera are 
known in Central America. 

Flowers in only 2-3 whorls; leaves deeply cordate at the base, the nerves very 
inconspicuous or almost obsolete; flowers polygamous (perfect and stami- 
nate) Lophotocarpus 

Flowers usually in numerous whorls; leaves attenuate to cordate at the base, con- 
spicuously nerved; flowers not polygamous, but either all perfect or else 
staminate and pistillate upon the same plant. 
Flowers either staminate or pistillate, both kinds on the same plant; leaves 

attenuate at the base or sagittate Sagittaria 

Flowers all perfect; leaves acute or cordate at the base, never sagittate. 

Echinodorus 

ECHINODORUS L. Richard 

Reference: N. C. Fassett, Echinodorus in the American tropics. 
Rh. 57: 133-158. 1955. 

Perennial or annual herbs, growing in water or wet soil; leaves ovate-rounded 
to almost linear, attenuate to cordate at the base; inflorescence of few to many 



76 FIELDIANA: BOTANY, VOLUME 24 

whorls of flowers, often paniculate, the flowers sessile or pedunculate, perfect; sepals 
herbaceous, usually coriaceous, costate-striate, persistent, after anthesis recurved, 
spreading, or appressed, sometimes accrescent; petals delicate, fugacious; stamens 
9 to many, the filaments linear, the anthers oblong, attached dorsally above the 
base, dehiscent by lateral slits; carpels usually numerous, 1-ovulate, forming a 
dense head, the style terminal; fruit heads mostly spheric or ovoid, sometimes 
echinulate, the fruits scarcely compressed, their sides costulate and sometimes 
glandular; seeds smooth or punctate. 

About 20 species are known, two in Europe and Africa, the others 
in temperate and tropical America. 

Carpels 20 or fewer in a loose head; stamens 6 or 9; anthers basifixed; stylar beak 

of nutlet 0.5 mm. or less long or obsolete. 
Nutlet with 3 smooth ridge-like facial ribs; blade of leaves linear-lanceolate to 

elliptic, tapered at base E. tenellus 

Nutlet with 1-several crested ribs; blade of emersed leaves ovate and deeply 

cordate E. nymphaeifolius 

Carpels many in a dense head; stamens 9-30; anthers versatile; nutlet with well- 
developed beak. 
Leaves with blades cordate at base, stellate-pubescent at base of blade and 

summit of petiole E. grandiflorus 

Leaves with blades tapered to truncate at base, glabrous E. Andrieuxi 

Echinodorus Andrieuxi (Hook. & Arn.) Small, N. Am. Fl. 17, 
pt. 1: 46. 1909. Alisma Andrieuxi Hook. & Arn. Bot. Beech. Voy. 
311. 1837. Echinodorus ellipticus 7 ovata Micheli in DC., Monogr. 
Phan. 3:52. 1881. 

Open or brushy marshes of the Oriente, 400-900 meters; Chiqui- 
mula; Jutiapa. Western Mexico; British Honduras; El Salvador; 
Nicaragua. 

A coarse stout perennial, sometimes a meter high but usually lower, with thick 
erect rhizomes, the scapes stria te and angulate; leaves few or numerous, on long, 
marginate or winged petioles, the blades shorter than the petioles, lanceolate to 
elliptic, as much as 35 cm. long and 14 cm. wide but usually smaller, acute or short- 
acuminate, acute or cuneate at the base, with 3-5 or sometimes 5 elevated nerves, 
not punctate; inflorescence simply spicate or spicate-paniculate, usually large, the 
whorls of flowers usually numerous, separated, the bracts lanceolate, long-acumi- 
nate; peduncles stout, 5-15 mm. long or sometimes even shorter; sepals broadly 
ovate, coriaceous, costate-striate, erect and somewhat enlarged in fruit; stamens 
20-24; fruit heads ovoid, mostly 8-10 mm. long, echinate; carpels rostrate, 6-8- 
costulate. 

Echinodorus grandiflorus (Cham. & Schlecht.) Micheli in DC. 
Monogr. Phan. 3: 57. 1881. E. grandiflorus y floribundus Micheli, 
loc. cit. Alisma grandiflorum Cham. & Schlecht. Linnaea 2: 132. 
1827. Figure 17. 




FIG. 17. Echinodorus grandiflorus. a, Inflorescence (X 
Leaf (X y%). 



. 6, Flower (X 4). 



77 



78 FIELD IANA: BOTANY, VOLUME 24 

In shallow water of swamps or at edges of lakes and streams, 600 
meters or less; Izabal; Chiquimula; Jutiapa; Santa Rosa. Honduras 
to Panama; Cuba; South America. 

A coarse perennial, often a meter high or more, with short thick rhizomes, the 
scapes angulate, sometimes sparsely muricate or pubescent; emersed leaves erect, 
long-petiolate (except in individuals from dry soil), densely pellucid-punctate, the 
blades ovate-rounded or broadly ovate, sometimes 50 cm. long and 40 cm. wide 
but usually smaller, 13-17-nerved, obtuse, at the base truncate to rather deeply 
cordate, with rounded basal lobes, glabrous or often more or less muricate or pubes- 
cent beneath on the nerves, the base of blade and summit of petiole stellate- 
pubescent; inflorescence large, simple or often much branched, the whorls of flowers 
few or numerous, remote, many-flowered, the peduncles long and slender, mostly 
1.5-4 cm. long; bracts ovate, long-acuminate; sepals ovate, obtuse, striate-costate, 
spreading or appressed in fruit, scarcely accrescent; fruit heads globose, about 1 cm. 
broad, obscurely echinate; nutlets winged on the back, with 3-4 ribs that are some- 
times slightly winged toward the summit, each side with usually 2 facial glands 
that are elongate and rounded at both ends and are placed well below the summit 
of the nutlet, and a short stout beak. 

Echinodorus nymphaeifolius (Griseb.) Buchenau, Bot. Jahrb. 
2: 483. 1882. Alisma nymphaeifolium Griseb. Cat. PL Cuba 218. 
1866. Helianthium nymphaeifolium Small, N. Amer. Fl. 17: 45. 1909. 

Collected at Maskall Pine Ridge, Gentle 1109, British Honduras, 
and in Campeche, and doubtless occurring in Pete"n. Cuba. 

Plants comparatively small, mostly 40 cm. tall or less; submersed leaves rib- 
bon-like, thin and flaccid, 15-20 cm. long, about 1 cm. wide; emersed leaves on 
very long petioles, the blades thin, ovate or oblong-ovate to rounded-ovate, 3-12 
cm. long, 1.5-9 cm. wide, 9-13-nerved, rounded at the apex, deeply and narrowly 
cordate at the base with scattered pellucid lines 0.2-0.8 mm. long; scapes solitary 
or several, the flowers forming a rather small but much branched erect ovate or 
conic panicle, the peduncles 1-2 cm. long, slender; sepals broadly ovate, 1-1.5 mm. 
long; petals scarcely as long as the sepals, suborbicular; fruit heads depressed, 
subtended by the persistent reflexed calyx; nutlets 1.4 mm. long, 1 mm. wide, with 
a beak 0.2 mm. long, flat-sided, with a broad crested keel, and with crested ribs 
and 1 or 2 long glands on each face, the ribs conspicuously cristate. 

Echinodorus tenellus (Mart.) Buch. var. latifolius (Seubert) 
Fassett, Rhodora 57: 202. 1955. Alisma tenellum forma latifolia 
Seubert in Mart. Fl. Bras. 3, pt. 1 : 195, pi. 13, fig. 2, nutlet and plant 
to left. 1848. 

Bogs, swamps, and wet meadows, at elevations mostly between 
500-1700 meters; Pete*n; Jalapa; Alta Verapaz. El Salvador and 
West Indies to Venezuela and southern Brazil. 

Submersed leaves linear to oblong-oblanceolate, 12-22 cm. long, thin and 
flaccid; emersed leaves smaller, rarely 10 cm. long, the blades only a few mm. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 79 

wide, linear-lanceolate to narrowly ovate; scape slender, 10 cm. or less tall, with 
the flowers in one or few whorls; flowers small; nutlet 1.4-1.8 mm. long, with 3 
smooth ridges on each face; beak 0.2-0.5 mm. long. 



LOPHOTOCARPUS Durand 

Perennial aquatic plants with long-petiolate, broad, deeply cordate leaves; 
scapes simple, the polygamous flowers borne in a few whorls of 2-3 long-peduncu- 
late flowers, the upper flowers staminate, the lower ones perfect; stamens 9-15, the 
filaments compressed; carpels numerous, densely inserted upon a convex receptacle, 
achenes crowded and compressed, cristate or winged, the fruit head more or less 
enveloped by the large accrescent sepals. 

As treated by Buchenau the genus consists only of the following 
species. North American authors usually refer to Lophotocarpus 
half a dozen plants of Mexico and the United States that Buchenau 
refers to Sagittaria, and which are conspicuously different in habit 
from L. guayanensis. 

Bogin (Mem. N. Y. Bot. Gard. 9: 192. 1955) places Lophotocarpus 
guayanensis under Sagittaria guayanensis, referring the Guatemalan 
material to subsp. guayanensis. 

Lophotocarpus guayanensis (HBK.) J. G. Smith, Kept. Mo. 
Bot. Gard. 6: 61. 1894. Sagittaria guayanensis HBK. Nov. Gen. & 
Sp. 1:250. 1816. 

In mud or shallow water, 1000 meters or less; Pete"n; Jutiapa; 
Santa Rosa. Mexico; Honduras; El Salvador; Panama; northern 
South America. 

Plants sometimes submerged or floating; petioles often much elongate, always 
much exceeding the blades, nodose-septate; blades broadly ovate to rounded, 5-7 
cm. long or shorter, obtuse to rounded and emarginate at the apex, deeply cordate 
at the base; scapes often shorter than the leaves, bearing only a few close whorls 
of large flowers; sepals broadly ovate, green, in fruit 8-11 mm. long; fruiting pedun- 
cles short and stout, often exceeded by the persistent bracts; fruit heads depressed- 
globose, 12-15 mm. broad; achenes obovate, 2-2.5 mm. long, short-rostrate, the 
sides tuberculate. 

SAGITTARIA L. 

Plants perennial, growing in shallow water or wet soil, sometimes submerged 
and floating (not in Central American species), arising from thick or tuberous rhi- 
zomes; leaves long-petiolate, various in form; scapes simple, at least below the 
inflorescence, the flowers monoecious or dioecious, in whorls of 3, the flowers of the 
upper whorls usually staminate, the lower pistillate; stamens few or many; carpels 



80 FIELDIANA: BOTANY, VOLUME 24 

very numerous, distinct, crowded upon a convex or globose receptacle; achenes 
numerous, compressed, rostrate. 

About 30 species in temperate and tropical regions of both hemi- 
spheres. In Central America only the following species are known. 

Leaves mostly lanceolate or linear-lanceolate, long-attenuate to the base. 

S. landfolia 
Leaves triangular-sagittate, with large, acute or acuminate basal lobes. 

S. latifolia 

Sagittaria lancifolia L. PI. Jam. Pug. 27. 1759. Figure 18. 

In shallow water or in muddy soil about lakes or along streams, 
600 meters or less; Alta Verapaz; Izabal; Jutiapa. Southern United 
States to Mexico and British Honduras; Honduras; Nicaragua; 
Costa Rica; Panama; West Indies; South America. 

Plants large and coarse, glabrous, often a meter high or taller; leaves borne on 
long thick spongy petioles, erect, the blades lance-linear to elliptic, 20-50 cm. long, 
mostly 2-8 cm. wide, acute or acuminate, acute to long-attenuate at the base, con- 
spicuously nerved; scapes simple or usually branched, the flowers on long slender 
spreading peduncles, the thin bracts lanceolate; corolla pure white, 2-4 cm. broad; 
fruit heads 1-1.5 cm. in diameter; achenes cuneate or obovate, short-rostrate, with 
a narrow dorsal wing. 

Bogin refers material from Guatemala, British Honduras, Costa 
Rica, Nicaragua, Panama, Mexico, and eastern United States to 
subsp. media (Micheli) Bogin (Mem. N. Y. Bot. Card. 9:214. 1955). 

Sagittaria latifolia Willd. Sp. PI. 4: 409. 1806. 

In marshes, at or little above sea level; Izabal. Southern Canada 
and United States to Mexico; Honduras; Nicaragua; Costa Rica; 
West Indies; South America. Hawaii (where introduced). 

Plants glabrous, usually 30-60 cm. tall; leaves long-petiolate, the blades nar- 
rowly or broadly triangular-sagittate, 10-40 cm. long, acute or acuminate, the large 
basal lobes acute to attenuate; scapes simple or branched, the peduncles 1-5 cm. 
long, slender, the bracts ovate; petals large and white; fruit heads 1.5-3 cm. in 
diameter; achenes very numerous, obliquely obovate, winged, with a horizontal 
apical beak. 

Called "arrow-head" in the United States, in reference to the 
shape of the leaves. The starchy rhizomes were used as food by 
many of the North American Indians. 



BUTOMACEAE 

Reference: Fr. Buchenau, Butomaceae, Pflanzenreich IV. 16. 1903. 




FIG. 18. Sagittana lancifolia. a, Habit (X %). b, Achene (X 27). c, 
Staminate flower (X 2^)- 

81 



82 FIELDIANA: BOTANY, VOLUME 24 

Perennial or perhaps sometimes annual herbs, growing in water or wet soil, the 
sap usually milky; leaves usually all basal, the stems often scapiform; flowers con- 
spicuous, regular, perfect; sepals 3, persistent, green; petals 3, generally larger than 
the sepals, mostly thin and fugacious; stamens free, hypogynous, either 9 or numer- 
ous and arranged in several verticels, the outer ones sterile; carpels of the ovary 
generally 6, sometimes numerous, the anthers 2-celled; ovules numerous in each 
cell, anatropous, inserted on the inner surface of the cell; style none or elongate; 
fruits as many as the carpels, free or more or less connate, the pericarp dry; seeds 
numerous, small, curved or straight; embryo hippo crepiform or straight; endo- 
sperm none. 

Four genera and about a dozen species, in tropical and temperate 
regions of both hemispheres. Only the following are represented in 
tropical America. 

Carpels of the ovary and fruit 6; stigma borne on a long style; leaves about as 
broad as long, rounded or cordate at the base Hydrocleis 

Carpels numerous; stigma sessile; leaves much longer than broad, attenuate at 
the base. . . . Limnocharis 



HYDROCLEIS L. Richard 

Herbs, usually growing in shallow water and with floating leaves, glabrous; 
leaves borne on very long petioles; scapes several or numerous, the flowers few or 
numerous, long-pedicellate, clustered at the end of the scape; sepals green, per- 
sistent; petals fugacious, yellow; stamens several or numerous, the outer ones with- 
out anthers, the filaments linear or lanceolate, the anthers basifixed, 2-celled, 
dehiscent by lateral longitudinal slits; carpels 3 or 6, lance-linear, subconnate at 
the base, attenuate into the style; fruits free, membranaceous, dehiscent along the 
inner side; seeds numerous, areolate-cristate. 

About 4 species, in tropical America. Only the following is known 
in North America. 

Hydrocleis Stand ley i Steyermark in Standl. & Steyerm. Field 
Mus. Bot. 23: 31. 1944. Figure 19. 

In shallow water, about 1000 meters; Jutiapa (type collected along 
railroad between Agua Blanca and Amatillo, Steyermark 30416}; So- 
lola (Finca Moca). Honduras; Guanacaste, Costa Rica. 

Stems numerous from a short rootstock; petioles terete, 25-30 cm. long; blades 
of the floating leaves oval or broadly oblong, 4-6.5 cm. long, 3-4 cm. wide, rounded 
at the base and apex or shallowly cordate at the base; scapes 17-29 cm. long, um- 
bellately 5-11-flowered, the bracts oblong-lanceolate or ovate-lanceolate, subacute 
or acute, 2-2.5 cm. long, the pedicels 3-5 cm. long; sepals lanceolate, subobtuse or 
subacute, 15-19 mm. long, 3.5-4 mm. wide, carinate dorsally; petals yellow, 6-7 
mm. long, oblong-obovate; fertile stamens 6, the sterile ones 2-3; ovaries 3; follicles 
membranaceous, 12-13 mm. long; seeds brown, 0.8 mm. long. 




FIG. 19. Hydrocleis Standleyi (X %). 
83 



84 FIELD IANA: BOTANY, VOLUME 24 

LIMNOCHARIS Humboldt & Bonpland 

Plants growing in or at the edge of water, glabrous; leaves usually numerous, 
erect or ascending; stems naked, scapiform, the flowers umbellate at the apex; 
sepals persistent and embracing the fruit head; petals fugacious; stamens numer- 
ous, the outer one sterile, the filaments complanate; anthers basifixed, 2-celled, 
dehiscent by longitudinal lateral slits; carpels 15-20, verticillate, laterally com- 
pressed; style none, the stigma sessile, extrorse; follicles verticillate, scarcely coher- 
ent, laterally compressed, semicircular, membranaceous, sulcate dorsally, dehiscent 
internally; seeds very numerous, curved, transversely multicostate. 

Two species, the other one described from Matto Grosso, Brazil. 

Limnocharis flava (L.) Buchenau, Abh. Nat. Ver. Bremen 2: 2. 
1868. Alisma flavum L. Sp. PI. 343. 1753. Cebolla de chucho. 

At edge of a small pool, about 850 meters; Jutiapa (near Jutiapa, 
Standley 74967). Chiapas; Panama; West Indies; South America; 
naturalized (?) in the East Indies. 

Plants perennial from a short thick erect rhizome, the scapes erect, 20-40 cm. 
high; leaves erect or ascending, not floating, often exceeding the scapes, long-petio- 
late, the petiole vaginate; leaf blades variable in shape, lanceolate to oblong-elliptic 
in Central American plants, sometimes broadly ovate in other regions, mostly 8-18 
cm. long, acute to rounded at the apex, attenuate at the base; inflorescences umbel- 
liform, 2-12-flowered, the pedicels 3-4 cm. long, somewhat dilated and trigonous 
above; flowers yellow, about 1.5 cm. broad, the sepals green, broadly ovate, obtuse; 
petals broadly ovate or suborbicular, longer than the sepals; mature follicles about 
1 cm. long. 

HYDROCHARITACEAE 

Reference: P. A. Rydberg, Elodeaceae, Hydrocharitaceae, N. 
Amer. Fl. 17: 67-74. 1909. 

Aquatic herbs, floating or submerged, the stems much abbreviated or some- 
times elongate and leafy; leaves linear or broader, sessile or petiolate; flowers regu- 
lar, unisexual or rarely perfect, solitary within a tubular 2-fid or 2-parted spathe or 
subtended by 2 complicate bracts, the staminate sometimes more than 1 in the 
spathe; spathes sometimes borne on elongate scapes and emersed; calyx of 3 herba- 
ceous or petaloid sepals; corolla of 3 thin petals, or wanting; stamens 3-12, the 
filaments distinct or united, the anthers linear, 2-celled; carpels of the ovary 3-15, 
united, the ovary 1-celled with several parietal placentae or 2-15-celled, the ovules 
numerous; styles or stigmas as many as the ovary cells; fruit baccate or utricular. 

About 14 genera, in tropical and temperate regions. The genus 
Elodea has been found in El Salvador and may well occur in Gua- 
temala. 

Leaves petiolate, the blades ovate or rounded Limnobium 

Leaves sessile, linear or ligular. 

Fruit 1-celled, indehiscent; plants of fresh water Vallisneria 

Fruit 6-12-celled, dehiscent by valves; plants of salt water Thalassia 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 85 

LIMNOBIUM L. Richard 

Plants usually floating, stoloniferous, the roots pendent in the water; leaves 
basal, petiolate, spongy-reticulate beneath; flowers dioecious or monoecious; stami- 
nate spathe pedunculate, bearing 2-3 long-pedicellate flowers; sepals 3, oblong; 
petals 3; stamens 6-12, distinct or united below, the anthers linear; pistillate spathe 
of 2 bracts, bearing a single short-pedicellate flower; ovary 6-9-celled; stigmas as 
many as the cells, 2-parted; fruit baccate, 1-9-celled, many-seeded. 

The genus contains 3 or 4 other species, all American. 

Limnobium stoloniferum (G. F. W. Mey.) Griseb. Fl. Brit. 
W. Ind. 506. 1861. Hydromistria stolonifera G. F. W. Mey. Fl. 
Esseq. 153. 1818. 

Floating in lake, about 500 meters; Jutiapa (Lago de Guija, 
Steyermark 31806). West Indies; South America. 

Plants producing abundant elongate stolons; leaves few or numerous, floating 
on the surface of the water and rosulate, the petioles spongy, 11-15 cm. long; leaf 
blades oblong-oval to rounded-oval, 4-6 cm. long, obtuse or rounded at the apex, 
acute at the base, entire; staminate sepals lanceolate, spreading; petals white, erect 
or spreading, linear-filiform, half as long as the sepals; filaments subulate, shorter 
than the anthers. 

THALASSIA Solander 

Marine herbs with elongate rootstocks; leaves clustered at the nodes of the 
rootstock, linear; scapes arising from the leaf clusters, the spathes tubular, 2-cleft; 
flowers dioecious, solitary; staminate flowers pedicellate, the perianth of 3 petaloid 
sepals; stamens 9, the filaments very short or obsolete, the anthers dehiscent later- 
ally; pistillate flowers almost sessile in the spathe; ovary 6-12-celled; fruit stipitate, 
mammillate or echinate, dehiscent by valves. 

One other species is known, in the Red Sea and Indian Ocean. 

Thalassia testudinum Konig in Konig & Sims, Ann. Bot. 2: 
96. 1805. 

In salt water, Izabal; noted only about Puerto Barrios. Florida; 
Honduras; West Indies; northern South America. 

Plants submersed, the base of the short stem covered by persistent remains of 
old leaves; leaves 2-5, sheathing at the base, linear-ligulate, mostly 40 cm. long or 
less, 6.5-11 mm. wide, rounded at the apex; scapes solitary; lobes of the spathe 
elliptic, papillose-dentate on the margins; perianth lobes oblong, 10-12 mm. long, 
rounded at the apex; anthers linear, 8 mm. long; stigmas 9-12, linear-fusiform, 
pilose, 1 cm. long; fruit ellipsoid-fusiform, short-stipitate, short-rostrate, mam- 
millate. 

This and other phanerogamic plants of salt water are seldom col- 
lected and little herbarium material of them is available. They 



86 FIELDIANA: BOTANY, VOLUME 24 

probably are much more abundant and common than one might in- 
fer from the specimens found in herbaria. They often are cast upon 
the beach after storms. 



VALLISNERIA L. Tape-grass 

Submerged perennial plants of fresh water, acaulescent or nearly so, with stolo- 
niferous rootstocks; leaves all basal, linear; staminate flowers numerous, crowded 
on a spadix, enclosed in a 2-3-parted spathe, at maturity detached and floating on 
the surface of the water; perianth of 3 sepals; stamens 1-3, usually 2; pistillate 
flowers solitary at the end of a long slender scape, floating on the surface of the 
water, subtended by a tubular 2-cleft spathe; sepals and petals each 3; ovary 
1-celled, with 3 parietal placentae; stigmas 3- and 2-lobate; ovules numerous, ortho- 
tropous; fruit cylindric, indehiscent. 

Probably two species, one American, the other European and 
Asiatic. 

Vallisneria americana Michx. Fl. Bor. Am. 2: 220. 1803. 
Figure 20. 

In fresh-water streams or lakes, at or little above sea level; Pete"n; 
Izabal. Temperate North America; Honduras. 

Leaves thin, 10-50 cm. long, 3-8 mm. wide, obtuse or subacute, usually 
minutely denticulate toward the apex; staminate peduncles 1-5 cm. long, the 
spathe ovoid, 8-10 cm. long; pistillate peduncles thread-like, 30-60 cm. long, in 
age spirally twisting and drawing the fruit under water; spathe tubular, 1-1.5 cm. 
long; hypanthium in flower 1-2 cm. long, in fruit 5-12 cm. long; sepals elliptic, 
2-3 mm. long; petals minute; stigmas sessile, 2-cleft. 

Well known in the United States under the name "wild celery." 
The plants are a favorite food of at least some kinds of wild ducks, 
and are supposed to impart a particularly good flavor to the flesh of 
birds feeding upon them. The canvasback ducks of Chesapeake Bay, 
Maryland, have long been famous among epicures on this account. 

The American material differs from the Old World V. spiralis L. 
chiefly in the staminate inflorescence, the staminate spathe of the 
American plants being shorter and thicker than that of the European 
and Asiatic plants. Possibly the American material should be treated 
as a variety of the Old World species, in which case the name would 
be V. spiralis var. americana (Michx.) Torr. 

TRIURIDACEAE 

Reference: Hanns Giesen, Triuridaceae, Pflanzenreich IV. 18. 
1938. 




n n 

X 50). c, Pistillate flower (X 3). 



- a ' Habit (X M- 6 Staminate flower 



87 



88 FIELDIANA: BOTANY, VOLUME 24 

Small, simple or branched, saprophytic herbs, without green coloring; leaves 
reduced to scales; flowers mostly minute, monoecious, dioecious, or perfect, the 
inflorescence racemose or sympodial; perianth 3-10-parted, the segments equal or 
alternately unequal, often caudate or barbate at the apex, valvate in bud; stamens 
2, 3, 4, or 6 in the staminate flower, inserted on a flat or convex receptacle, the fila- 
ments short and connate at the base or none; anthers 2-4-celled, the cells finally 
confluent, dehiscent by longitudinal or transverse slits; pistillate flower without 
stamens or staminodia, the carpels of the ovary numerous, inserted on a more or 
less convex receptacle, smooth or papillose; mature carpels usually vertically de- 
hiscent; style terminal, lateral, or basal, subulate or clavelliform, sometimes penicil- 
late-papillose at the apex; seed ellipsoid; perfect flowers with a variable number of 
stamens, the filaments short, not connate, the anthers 3-4-celled, the carpels 
numerous, like those of the pistillate flower. 

Seven genera and about 80 species, in the tropics of both hemi- 
spheres. Only the following genera are represented in Central 
America. 

Perianth segments 4 Soridium 

Perianth segments of a different number from 4 in American species. 

Perianth segments long-caudate; flowers dioecious Triuris 

Perianth segments not caudate; flowers monoecious Sdaphila 



SCIAPHILA Blume 

Roots usually pilose, slender; stems slender, erect, irregularly flexuous, gla- 
brous; inflorescence racemose, the pedicels alternate; flowers monoecious, perfect 
or unisexual; perianth segments 6 in the Guatemalan species; stamens 6 in the 
local species; anthers 3-4-celled, the connective not produced; carpels of the ovary 
numerous, the style penicillate, papillose, or glabrous at the apex. 

Species about 45, in the tropics of both hemispheres. One other 
Central American species is known from Panama. In spite of their 
usual scarcity locally, and the fact that the plants grow in sheltered 
places from which one would suppose dispersal of the minute seeds 
would be difficult, some of the species of this and other genera seem 
to have a very wide distribution. 

Sciaphila picta Miers, Trans. Linn. Soc. 21 : 48. 1852. Figure 21. 

Wet dense forest, usually among rotting leaves, at or near sea 
level; Izabal. Atlantic coast of Honduras; Panama; Colombia. 

Plants mostly 8-15 cm. high, coral-red, or the flowers dark red; leaves bract- 
like, ovate and clasping, 2-2.5 mm. long; racemes 4-13 cm. long, many-flowered, 
secund, laxly 10-30-flowered; bracts about 1.5 mm. long, ovate, acute; pedicels 
2-3 mm. long, decurved; staminate flowers none; perfect flowers 2-2.5 mm. broad; 
perianth segments 6, equal, ovate-lanceolate, subobtuse, obscurely barbate at the 
apex; stamens 6, opposite the perianth segments, the filaments very short; carpels 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 89 




FIG. 21. Sciaphila picta. a, Habit (X ]4)- b, Stamen (X 117). c, Flower, 
from above (X 23). d, Flower, lateral view (X 23). e, Pistil (X 117). 



of the ovary 10-15, the style lateral, penicillate-papillose at the apex, longer than 
the carpel. 

SORIDIUM Miers 

Small slender erect plants with more or less flexuous stems, the flowers race- 
mose, alternate; perianth segments 4; stamens 2 in the staminate flower, opposite 

2 opposing perianth segments, sessile or nearly so; anthers 2-celled, dehiscent by a 
transverse and an apical slit; carpels of the ovary numerous in the pistillate flower, 
the style lateral, smooth and obtuse or sometimes penicillate-papillose at the apex; 
fruit indehiscent. 

Two species are known, the other in British Guiana. 

Soridium Spruceanum Miers, Trans. Linn. Soc. 21: 50. 1852. 

Dense wet forest, usually among rotting leaves, at or little above 
sea level; Izabal (along Rio Frlo, Steyermark 39899, 39897). British 
Honduras; Venezuela, Guianas, and northern Brazil. 

Plants slender, simple, 5-25 cm. high, white throughout; leaves bract-like, 

3 mm. long, narrowly lanceolate, acute; racemes 1-8 cm. long, laxly 10-30-flow- 
ered, not secund; bracts 2.5-4 mm. long, narrowly lanceolate; pedicels 1.5-2.5 mm. 



90 FIELDIANA: BOTANY, VOLUME 24 

long, erect-spreading, not decurved; staminate flower 2 mm. broad, the segments 
equal, ovate, very minutely barbate at the apex, minutely papillose within; pistil- 
late flower 2 mm. broad, the perianth segments like those of the staminate flower; 
carpels of the ovary about 20, verruculose above; style slightly longer than the 
carpels. 

This has been reported from British Honduras as Sciaphila picta 
Miers. 

TRIURIS Miers 
The genus consists of only the following species: 

Triuris hyalina Miers, Trans. Linn. Soc. 19: 77. 1845. T. brevi- 
stylis Bonn. Smith, Bot. Gaz. 16: 14. 1891 (type from Pansamald, 
Alta Verapaz, 1,200 meters, Tuerckheim 1884). 

Known in Guatemala only from the collection cited. Brazil. 

Root slender, sparsely pilose, simple; stems 2.5-7.5 cm. high, simple or branched 
from the base, leafless; inflorescence sympodial; bracts lanceolate, acute, auriculate 
at the base; flowers dioecious; perianth segments of the staminate flower 3, ovate- 
triangular, contracted at the apex into a caudicle as much as 9 mm. long, the blade 
5 mm. long; stamens 3, alternate with the perianth segments, immersed near the 
base of the rounded torus, sessile, the anthers 4-celled, dehiscent by longitudinal 
slits; pistillate perianth like that of the staminate flower; carpels very numerous, 
the style terminal, smooth, glabrous, longer than the carpel. 

CYPERACEAE. Sedge Family 

References: Standley, The Cyperaceae of Central America, Field 
Mus. Bot. 8: 239-292. 1931; Hugh T. O'Neill, The sedges of the 
Yucatan Peninsula, Carnegie Inst. Wash. Publ. 522: 249-322. 1940. 

Grass-like or rush-like plants, herbaceous, annual or perennial, often with rhi- 
zomes; stems (culms) slender, solid, triangular, quadrangular, terete, or com- 
pressed; leaves narrow and grass-like, with closed sheaths; flowers perfect or 
imperfect, arranged in spikelets, one in the axil of each glume, the spikelets soli- 
tary or clustered, containing one to many flowers; scales 2-ranked or spirally im- 
bricate, persistent or deciduous; perianth hypogynous, composed of bristles or 
scales, rarely calyx-like, often wanting; stamens usually 1-3, the filaments slender 
or filiform, the anthers 2-celled; ovary 1-celled; ovule 1, erect; style 2-3-cleft or 
rarely simple; fruit an achene, lenticular, plano-convex, or trigonous. 

One of the largest families of plants, distributed through almost 
all parts of the earth. The largest genus, Carex, is most abundantly 
represented in temperate and arctic regions, but many of the other 
large groups, such as Cyperus and Scleria, are best developed in the 
tropics. At least five additional genera are represented in other parts 
of Central America, and one or two of them might be found in Gua- 
temala, although this is doubtful. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 91 

Pistillate flowers enclosed in a more or less inflated utricle. 

Utricle enclosing a long uncinate bristle Uncinia 

Utricle enclosing only the pistil Carex 

Pistillate flowers not enclosed in a utricle, merely subtended by glumes. 
Culms bearing a single spikelet. 

Leaves all reduced to bladeless sheaths Eleocharis 

Blade-bearing leaves present. 

Scales of the spikelets spirally imbricate; base of the style persistent. 

Bulbostylis 
Scales of the spikelets 2-ranked; base of the style not persistent. 

Abildgaardia 

Culms bearing more than a single spikelet. 
Scales of the spikelets evidently 2-ranked; perianth none. 

Spikelets with only one perfect flower, capitate Kyllinga 

Spikelets usually with 2 to many perfect flowers, if 1-flowered then not 

capitate Cyperus 

Scales spirally imbricate; a perianth of bristles often present. 
Flowers all unisexual. 

Fertile flower in each spikelet one, naked, terminal Hypolytrum 

Fertile flowers in each spikelet often more than one, each subtended by 
a scale, lateral. 

Fertile flower basal Scleria 

Fertile flower terminal Calyptrocarya 

Flowers at least partly (the fertile ones) perfect. 
Empty scales at the base of the spikelet 3 or more. 

Style 3-cleft Cladium 

Style 2-cleft. 

Spikelets somewhat compressed, clustered in a single head; bristles 
none Dichromena 

Spikelets not compressed, usually paniculate or corymbose; bristles 
usually present Rhynchospora 

Empty scales at the base of the style usually only 1-2. 

Base of the style persistent upon the achene as a tubercle . . Bulbostylis 
Base of the style not persistent. 

Flowers without inner perianth scales. 

Base of the style swollen; bristles none Fimbristylis 

Base of the style not swollen; bristles present Scirpus 

Flowers with one or more inner perianth scales. 

Flowers with 3 broad stipitate scales alternating with bristles or 
perianth of 6 bristles; plants perennial, with broad leaves. 

Fuirena 

Flowers with 1-2 hyaline scales and no bristles; annuals with 
narrow leaves. 

Inner scales 2, convolute Lipocarpha 

Inner scale 1, minute Hemicarpha 



92 FIELDIANA: BOTANY, VOLUME 24 

ABILDGAARDIA Vahl 

Glabrous perennials with slender culms, the narrow, setaceous basal leaves 
flattened; spikelets usually solitary, subtended by an involucre of one bract; scales 
imbricated in two rows, keeled and deciduous; bristles none; stamens 1-3; style 
pubescent, deciduous, with a somewhat swollen base; achenes trigonous. 

About 15 species, chiefly in the Old World tropics. Only the fol- 
lowing species is known from Central America. 

Abildgaardia monostachya (L.) Vahl, Enum. 2: 296. 1805. 
Cyperus monostachyos L. Mant. 180. 1771. Fimbristylis monostachya 
Hassk. PI. Jav. Rar. 61. 1848. Covolillo. 

Savannas or dry slopes, 1600 meters or less; Pete"n; Huehueten- 
ango. Florida; Yucatan and Tabasco to Panama; South America; 
Old World tropics. 

A cespitose perennial, the base of the plant usually hard and bulb-like, the 
culms wiry, often twisted, 10-40 cm. tall, glabrous; leaves setaceous, 10-30 cm. 
long, 0.5-1.5 mm. wide, about half as high as the culms, flat or involute, scaberu- 
lous on the margins; involucral bracts 1 or 2, shorter than the spikelet; spikelet 
ovate-lanceolate, 20-40-flowered, 10-20 mm. long, 5 mm. wide; scales ovate, acute, 
cuspidate, chartaceous, closely imbricate, 5-7 mm. long, strongly keeled; style with 
3 short branches; achene depressed-globose, umbonate, broadly stipitate, 1.5-2.5 
mm. long, yellowish-white or brown, conspicuously verrucose or muricate. 



BULBOSTYLIS 

Reference: Henry K. Svenson, Scirpeae, N. Amer. Fl. 18: 540- 
. figs. 8-21. 1957. 

Annuals or perennials, the culms slender, sometimes filiform, leafy below, the 
leaves narrowly linear or filiform, the sheaths ciliate or pubescent; spikelets um- 
bellate, capitate, or solitary, subtended by one to several bracts, the scales spirally 
imbricate, commonly deciduous, mostly pubescent; flowers perfect; perianth none; 
stamens 2-3; style 2-3-cleft, glabrous, its base enlarged and persistent as a tubercle 
at the apex of the achene; achene 3-angulate, turgid or lenticular. 

About 70 species, in temperate and warm regions of both hemi- 
spheres. All but one of the known Central American species are 
listed here. 

Spikelets solitary at the apex of the culm, sometimes also borne at the bases of 
the culms. 

Spikelets acuminate B. Funckii 

Spikelets ovoid, obtuse. 

Plant with a thickened woody base B. paradoxa 

Plant not woody-thickened at the base B. pubescens 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 93 

Spikelets capitate or umbellate at the apex of the culm, always more than one. 
Spikelets all sessile or nearly so, in head-like clusters, these solitary or umbellate. 

Plants slender annuals; achenes 0.6 mm. long B. barbata 

Plants perennials; achenes 0.7-0.9 mm. long. 

Plants glabrous B. junciformis 

Plants copiously pubescent, at least on the culms and leaves B. vestita 

Spikelets all or mostly pedicellate, often solitary on the rays of the umbel. 
Achenes (when mature) prominently spinescent-papillose, 0.6-0.8 mm. long; 
pedicels delicate, mostly arcuate-upcurved; spikelets narrowly lanceolate. 

B. tenuifolia 

Achenes (when mature) not prominently spinescent-papillose, commonly 

transversely rugose or with wavy lines, 0.75-1.2 mm. long; pedicels 

thicker and firmer, mostly straight; spikelets lance-ovate to ovoid-oblong. 

Mature achenes stramineous; slender annuals; achenes 0.75-0.9 mm. long. 

B. capillaris 

Mature achenes dark gray; annuals or perennials; achenes 0.8-1.2 mm. long. 

Slender annuals B. arenaria 

Perennials with hard thickened base B. juncoides 

Bulbostylis arenaria (Nees) Lindm. Bih. Sv. Vet.-Akad. Handl. 
26 (3) 9: 19. 1900. Isolepis arenaria Nees, Linnaea 9: 291. 1834, 
nomen nudum. Oncostylis arenaria Nees in Mart. Fl. Bras. 2, pt. 1: 
87. 1842. 

At 100 meters or less. Sandy pinelands, British Honduras 
(Schipp 919); West Indies; northeastern South America to eastern 
Brazil. 

Plants annual, tufted, the culms slender, 5-25 cm. high, glabrous or rarely 
minutely strigulose; leaves half as long as the culms or shorter; spikelets 2-6 mm. 
long, acute; scales brown, keeled, acute to cuspidate; achenes obovate, 0.8-1.1 mm. 
long, transversely rugose, becoming dark gray when mature. 

The specimen from British Honduras, determined by Dr. Sven- 
son, has not been examined by the authors. 

Bulbostylis barbata (Rottb.) Clarke in Hook. f. Fl. Brit. Ind. 
6: 651. 1893. Scirpus barbatiLS Rottb. Descr. PI. Rar. 27. 1772. 
S. barbatus Rottb. Descr. 52. pi. 17, j. If. 1773. 

At 200 meters; Solola (near the railroad station at Guatalon, 
Steyermark 48075). Southeastern United States; Martinique; South 
America; widely distributed in the Old World tropics. 

Plants annual, densely tufted, the culms very slender, 5-20 cm. high, glabrous 
or nearly so; leaves half as long as the culms, setaceous, almost glabrous, the sheaths 
bearing numerous long white hairs at the apex; spikelets 3-12, sessile in a dense 
head or compact umbel 1-1.5 cm. in diameter, ferruginous to dark brown, 3-10 mm. 



94 



FIELDIANA: BOTANY, VOLUME 24 



long, linear to oblong-lanceolate, acute, 3-15-flowered; scales rather distant on the 
rachilla, cymbiform, ovate-lanceolate, strongly keeled with a green midrib and 
brown sides and a small subrecurved mucro, mostly sparsely strigose-hispidulous 
or rarely glabrous; style equaling the achene, the persistent base minute, red or 
blackish; achene trigonous, obovate or obovoid, 0.6 mm. long, pale or dull brown, 
subtruncate, smooth or obscurely reticulate. 

It would be interesting to know in what manner this plant, mostly 
of Asia and Africa, has reached the lowlands of Guatemala. 




FIG. 22. Bulbostylis capillaris. a, Habit (X Y$). b, Inflorescence (X 
c, Achene (X 42). 



Bulbostylis capillaris (L.) Clarke in Hook. f. Fl. Brit. Ind. 6: 
652. 1893. Scirpus capillaris L. Sp. PL 49. 1753. Fimbristylis capil- 
laris Gray, Man. 530. 1848. Stenophyllus capillaris Britton, Bull. 
Torrey Club 21: 30. 1894. Figure 22. 

Sandy stream beds and rocky slopes, often in pine-oak forest, 
250-1900 meters; Zacapa; Quezaltenango; Huehuetenango. United 
States to Mexico; British Honduras; Honduras; Panama; Cuba. 

Plants annual, densely cespitose, the culms very slender, 5-30 cm. tall, 4-6- 
striate, glabrous or minutely scaberulous at the apex; leaves 3-15 cm. long, about 
1 mm. wide, filiform, shorter than the culms, bearing a tuft of very long hairs at 
the apex of the sheath, scabrous on the margins, otherwise usually glabrous or 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 95 

nearly so; bracts 1-3, setaceous, short; spikelets ovoid-oblong, 5-8 mm. long, about 
1.5 mm. wide, mostly 6-14-flowered, long-pedicellate, in simple or compound um- 
bels; scales ovate, rounded to emarginate at the apex, dark brown, the keel green, 
minutely puberulent or fimbriate; achene trigonous, obovate or obovoid, 0.75-0.9 
mm. long, stramineous to pale brown, truncate at the apex with a small conic 
tubercle. 

Bulbostylis Funckii (Steud.) Clarke in Urban, Symb. Antill. 
5: 290. 1907. B. Funckii (Steud.) Clarke, Kew Bull. Add. Ser. 8: 26. 
1908. Isolepis Funckii Steud. Syn. PL Glum. 2: 91. 1855. Steno- 
phyllus Funckii Britton, Bull. Torrey Club 21: 30. 1894. 

Damp meadows, about 1800 meters, or lower; Huehuetenango 
(along road east of San Sebastian, Standley 81476); reported also 
from San Marcos (Malacatan). Southwestern United States; Mex- 
ico; El Salvador; South America. 

A dwarf annual, the filiform culms and leaves 3-12 cm. high, the culms striate, 
glabrous or nearly so, the very narrow leaves scaberulous on the margins; spikelets 
linear to lance-oblong, acuminate, solitary at the apex of the culm and densely 
clustered and sessile at the base of the plant in the leaf axils, 4-10 mm. long; basal 
spikelets few-flowered and sometimes reduced to a single flower; scales of the ter- 
minal spikelet lanceolate, acute to mucronate, dark brown or castaneous with green 
keel, loosely attached, the lowest one often elongated and bract-like; achene trigo- 
nous, broadly obovate, 1-1.5 mm. long, white or stramineous to gray, obscurely to 
deeply transverse-rugulose, tubercle narrowly to broadly conic. 

A curious plant because of its unusual mode of inflorescence, not 
matched in any other Central American member of this family (ex- 
cept very rarely in one species of Eleocharis) . 

Bulbostylis junciformis (HBK.) Lindman, K. Sv. Vet.-Akad. 
Handl. 26 (3) 9: 19. 1900. Isolepis junciformis HBK. Nov. Gen. & 
Sp. 1: 222. 1815. Stenophyllus junciformis Britton, Bull. Torrey 
Club 43: 442. 1916. 

Rocky hillsides or in open grassy places, sometimes in pine forest, 
200-1800 meters; Zacapa; Jutiapa; Huehuetenango. British Hon- 
duras; El Salvador; Honduras; Costa Rica; Panama; Cuba; south- 
ward in South America to Brazil. 

A glabrous perennial with bulbous-thickened culm bases and fibrous roots, the 
culms cespitose, filiform, rigid, striate, 15-45 cm. tall; leaves 6-30 cm. long, shorter 
than the culms, frequently glaucous, sometimes scaberulous on the margins, the 
sheaths dull castaneous to dark brown, the apices long-fibrillose; bracts often 
shorter than the inflorescence, this consisting of a compound umbel, open or usually 
condensed, narrowly umbellate, the capitate glomerules on rays 1-3 cm. long or 
reduced to a single capitate inflorescence; spikelets fasciculate in clusters of 3-5, 
lanceolate, acute, commonly 4-8 mm. long, 3-5-flowered; scales lanceolate, acute 



96 FIELDIANA: BOTANY, VOLUME 24 

to short mucronate with the excurrent keel, the sides castaneous to reddish brown, 
minutely pubescent; achene bluntly trigonous, obovate or obovoid-cuneate, 0.7- 
0.9 mm. long, white or pale stramineous, when mature becoming bluish-gray, 
papillate, tipped with a small black tubercle. 

Bulbostylis juncoides (Vahl) Kukenth. in Osten, Anal. Mus. 
Hist. Nat. Montevideo II. 3: 187. 1931. Schoenus juncoides Vahl, 
Enum. 2: 211. 1805. Bulbostylis hirtella sensu Clarke, in Urban, 
Symb. Ant. 2: 166. 1900, not Isolepis hirtella Schrad. in Schult. 
Mant. 2: 70. 1824. Bulbostylis Langsdorffiana Clarke, in Urban, 
Symb. Ant. 2: 89. 1900, as to Mexican specimens determined by 
Clarke in Kew herb. Fimbristylis capillaris var. pilosa Britton, Bull. 
Torrey Club 15: 102. 1888 (type from Santa Rosa, Baja Verapaz, 
Tuerckheim 1283 [1293 as cited by Svenson in N. Amer. Fl. 18: 550. 
1957]). Zacatillo (Pete"n). 

Oak forest, dry open slopes, cornfields, and sandbars, mostly at 
1500-2500 meters; reported from Pete"n; Baja Verapaz; Chimal- 
tenango; Quezaltenango; Huehuetenango. Southwestern United 
States; Mexico to Honduras; West Indies; South America. 

Plants perennial with hard, thickened, somewhat bulbous base and coarse 
roots, densely cespitose, the culms filiform, sulcate, mostly 10-15 cm. high, some- 
times as much as 30 cm., glabrous to pilose; leaves setaceous, erect, often glaucous, 
usually one-third the length of the culms, glabrous to short-pilose; sheaths dark 
brown to cinnamon, fimbriate at the apex; inflorescence in the typical plant con- 
gested and not radiate, but more often open and umbellate with rays 0.5-2 cm. 
long bearing single spikelets; spikelets lance-ovate, subacute, 3-8 mm. long, casta- 
neous; scales broadly ovate, obtuse to acute, green-carinate, the keel short-ex cur- 
rent in the lowest glumes, frequently glutinous, glabrous to hirtellous; achene 
trigonous, obovoid or broadly obovate, 0.8-1.2 mm. long, yellowish brown or when 
mature dark gray, when mature rugulose with transverse wavy lines, tipped with 
a rounded-conic, dark brown, sessile tubercle. 

Most of the Guatemalan material may be referred to var. am- 
pliceps Kukenth. , in which the inflorescence is radiate with the rays 
of the umbels bearing single spikelets. 

Bulbostylis paradoxa (Spreng.) Lindm. Bih. Sv. Vet.-Akad. 
Handl. 26 (3) 9: 17. 1900. Schoenus paradoxus Spreng. Syst. Veg. 
1: 190. 1825. Schoenus spadiceus HBK. Nov. Gen. & Sp. 1: 227. 
pi. 69, f. 1. 1815, not Schoenus spadiceus Vahl, 1805. B. paradoxa 
Clarke, Kew Bull. Add. Ser. 8: 109. 1908. Stenophyllus paradoxus 
Standl. Contr. U. S. Nat. Herb. 18: 88. 1916. B. spadicea (HBK.) 
Kukenth. Repert. Sp. Nov. 23: 197. 1926. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 97 

British Honduras (El Cayo District, sandy pine uplands), and 
probably extending into Pete"n. Oaxaca; Cuba; Costa Rica; Pan- 
ama; southward in South America to Brazil. 

A glabrous perennial, usually forming very large and dense, hard clumps, aris- 
ing from a thick caudex often 4-5 cm. in diameter; culms 4-12, very slender, striate, 
5-20 cm. tall, smooth; leaves numerous, filiform and wiry, equaling or shorter than 
the culms, less than 0.5 mm. wide, ciliate, the short densely lanate sheaths with 
long hairs at the apex; bracts absent; spikelets solitary, obovate, 6-10 mm. long, 
3-5 mm. wide; scales lance-subulate, rigid, 3-nerved, lanate-fimbriate on the keel, 
this excurrent as a mucro 2 mm. long; achene trigonous, obovate to pyriform, 1.5 
mm. long, pale to dark brown, transversely rugose. 

This is typically a savanna plant which evidently is exceptionally 
resistant to fires to which such areas generally are subjected in Cen- 
tral and South America. At the base of the hard mass of green leaves 
and culms there usually is a dense coat consisting of the bases of the 
leaves of former years, badly scorched and charred. 

Bulbostylis pubescens (Presl) Svenson, N. Amer. Fl. 18: 542. 
1957. Abildgaardia pubescens Presl, Rel. Haenk. 1: 180. 1828. Fim- 
bristylis Preslii Kunth, Enum. PI. 2: 228. 1837. 

At 200 meters; Zacapa (dry plains, near Estanzuela, Steyermark 
29089). Mexico; El Salvador; Costa Rica to Colombia. 

Plants annual or perhaps perennial, the culms densely cespitose, wiry and slen- 
der, striate, 10-30 cm. high, about 0.6 mm. thick at the apex, glabrous to slightly 
hispid, minutely somewhat roughened; leaves setaceous, much shorter than the 
culms, usually less than half as long as the culms, glabrous to pilose with short 
spreading hairs, the sheaths stramineous, long-pilose at the apex; spikelets solitary 
at the apex of the culm, brown, ovoid to lance-oblong, 5-13 mm. long, 3-6 mm. 
wide; scales broadly ovate, acute to obtuse, sometimes mucronate, stramineous to 
black, little or not at all keeled with greenish midrib, nearly glabrous to hirtellous; 
achene trigonous, obpyramidal to broadly obovate, truncate to depressed at the 
apex and deeply 3-lobed, abruptly narrowed below the middle to a stipitate base, 
1.2-1.3 mm. long, pale brown or yellowish, prominently transversely rugose when 
mature, tipped with a conic trigonous tubercle. 

Bulbostylis tenuifolia (Rudge) Macbr. Field Mus. Bot. 11: 5. 
1931. Scirpus tenuifolius Rudge, PI. Guian. 18. 1805. Bulbostylis 
capillaris var. tenuifolia Clarke, Symb. Ant. 2: 89. 1900. 

At 2000 meters or lower; Guatemala; Jalapa; Huehuetenango. 
Mexico; El Salvador; Honduras; Costa Rica; Panama; Cuba; north- 
ern South America. 

Plants chiefly annual, densely cespitose, the culms slender, 5-30 cm. tall, stri- 
ate, glabrous or rarely hirtellous; leaves 3-15 cm. long, filiform, shorter than the 
culms, mostly glabrous; inflorescence usually broader than high in a simple or 



98 FIELDIANA: BOTANY, VOLUME 24 

compound umbel; spikelets narrowly lanceolate, acute, 3-6 mm. long, solitary and 
mostly long-pedicellate at the ends of usually upwardly curved arcuate delicate 
pedicels; scales dark brown, obtuse to acute, glabrous to hirtellous; achene trigo- 
nous, obovate, 0.6-0.8 mm. long, stramineous to gray-brown, when mature finely 
spinescent-papillose, when immature almost smooth to somewhat reticulate, tipped 
by a minute reddish-brown tubercle. 

Bulbostylis vestita (Kimth) Clarke, Symb. Ant. 2: 87. 1900. 
Isolepis vestita Kunth, Enum. PI. 2: 210. 1837. Stenophyllus vestitus 
Britton, Bull. Torrey Club 43: 446. 1916. 

Probably in savannas, Pete"n (La Libertad). Southern Mexico 
and British Honduras; Honduras; West Indies; South America. 

Plants perennial, usually densely cespitose, sometimes with enlarged culm 
bases, the very slender culms rigid, 10-40 cm. tall, strigose-pubescent; leaves seta- 
ceous, 25 cm. long or less, usually shorter than the culms, densely strigose-pubes- 
cent, the sheaths stramineous to light brown, long-fimbriate at the apex; bracts 
very short; spikelets forming a single terminal head or a simple umbel of few heads, 
3-10 mm. long; scales ovate, coriaceous, reddish brown, frequently glutinous, 
hirtellous, the yellowish keel prominently mucronate, the mucro about 1 mm. 
long; achene trigonous, obovate or obo void-cuneiform, 0.8-0.9 mm. long, white to 
brown with a prominent yellow stripe beneath the blunt angles, when mature cel- 
lular-reticulate or minutely papillate, tipped with a minute, pale brown, flattened 
tubercle. 

CALYPTROCARYA Nees 

Cespitose perennials with more or less leafy culms and long narrow leaves; 
spikelets crowded in small dense heads, laxly corymbose-paniculate, the panicles 
terminal and axillary; spikelets unisexual, very small, fasciculate and sessile, each 
pedicel bearing a capitate cluster of false spikelets, each of these consisting of a 
terminal globose 1-flowered pistillate spikelet and several smaller ovoid 1-4-flow- 
ered staminate spikelets; stamen 1; bristles none; style slender, the branches 2-3, 
filiform; achene subglobose, not beaked. 

About 7 species in tropical America. Only one spec : es is known 
from Central America. 

Calyptrocarya glomerulata (Brongn.) Urban, Symb. Antill. 
2: 169. 1900. Becquerelia glomerulata Brongn. in Duperrey, Voy. 
Coquille 2: 163. 1829. C. fragifera Kunth, Enum. PL 2: 364. 1837. 
Figure 23. 

Densely forested swamps, at 350 meters or less; Alta Verapaz 
(Cubilgiiitz) ; Izabal. Chiapas to British Honduras and Panama, 
and in many regions of South America. 

Plants glabrous or nearly so, commonly 30-50 cm. high, with short rhizomes 
or none, the culms trigonous, almost covered by the sheaths, these purple and 
usually overlapping, sometimes puberulent; leaves 20-50 cm. long, 3-7 mm. wide, 




FIG. 23. Calyptrocarya glomerulata. a, Habit (X 1 A). b, Pistillate spikelet 
X 25). c, Cluster of spikelets (X 12). d, Achene (X 21). 



99 



100 FIELDIANA: BOTANY, VOLUME 24 

antrorsely scaberulous on the margins, 3-nerved; bracts large and similar to the 
leaves; inflorescence about three-fourths the length of the whole plant, with 4-6 
umbelliform panicles, the peduncles 1-2 cm. long, each bearing 6-10 pedicels; false 
spikelets 3-6, the pistillate spikelets 1.5 mm. long, the staminate spikelets 1-4- 
flowered; style branches 2; achene 1 mm. long and broad, lenticular, at first white 
and lustrous, becoming dull brown, puberulent, umbonate at the apex. 

Common about Puerto Barrios, where it seems to be confined to 
Manicaria swamps. 

CAREX L. 

Reference: Kennenth K. Mackenzie, Cariceae, N. Amer. Fl. 18: 
3-478. 1931-35; North American Cariceae, pis. 1-539. 1940. 

Plants grass-like, perennial by rhizomes, the culms mostly trigonous; leaves 
3-ranked, the upper ones (bracts) elongate or short and subtending the flower 
spikes, or wanting; flowers usually monoecious, sometimes dioecious, the flowers 
solitary in the axils of the glumes or scales, forming short and head-like or often 
much elongate spikes; spikes one to many, wholly pistillate, wholly staminate, 
androgynous (with staminate flowers above and pistillate ones below), or gynaecan- 
drous (with pistillate flowers above and staminate ones below); perianth none; 
stamens 3; pistillate flower of a single pistil, with a style and 2-3 stigmas; achene 
trigonous or lenticular, completely enveloped in a tight or inflated perigynium. 

One of the largest genera of plants, with more than 1000 species, 
found in most parts of the earth but most abundant in temperate or 
cold regions. Very few species are found in the tropics, and those 
chiefly in the mountains at higher elevations. About three other 
species have been found in Costa Rica and Panama. 

Spikes sessile, short and head-like; achenes lenticular, the stigmas 2; perigynia 

glabrous. 
Terminal or all the spikes androgynous. 

Rootstocks long-creeping; perigynia 1.5 mm. wide, spongy at base. 

C. praegracilis 
Rootstocks short-creeping; perigynia 2-3 mm. wide, not spongy at base. 

C. xalapensis 
Terminal or all the spikes gynaecandrous. 

Body of the perigynium widest above the middle; heads pale green . . C. Longii 
Body of the perigynium widest at or below the middle; heads mostly brownish 

or castaneous C. peucophila 

Spikes usually conspicuously pedunculate, if sessile the spikes much longer than 
broad; achenes lenticular and with 2 stigmas, or triangular in cross section 
and with 3 stigmas; perigynia glabrous or sometimes pubescent. 
Perigynia scabrous or pubescent. 

Body of the perigynium completely filled by the achene C. planostachys 

Body of the perigynium not completely filled by the achene, the upper part 
empty. 

Leaves 1-2.5 mm. wide; scales short-aristate at the apex C. scabrella 

Leaves 2.5-4 mm. wide; scales long-aristate . . . .C. polystachya var. Bartlettii 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 101 

Perigynia glabrous. 

Stigmas 2; perigynia abruptly contracted into a minute beak. 

C. cuchumatanensis 

Stigmas 3 ; perigynia with a conspicuous, often elongate beak, often attenuate 

to the beak. 
Spikes paniculate, the lowest peduncles of the inflorescence branched. 

C. polystachya 
Spikes not paniculate, the peduncles all simple, the spikes sometimes sessile 

or nearly so. 

Achenes continuous with the style, the style persistent; perigynia some- 
what inflated; spikes about 8 mm. thick C. Thurberi 

Achenes jointed with the style, the style withering in age; perigynia not 

at all inflated; spikes usually conspicuously less than 8 mm. thick. 
Scales of the spikes blackish, castaneous, or ferruginous, the spikes very 

dark-colored or ferruginous. 

Pistillate spikes mostly 4 cm. long or shorter; achenes not outcurved. 
Scales dark brown-purple to blackish; spikes all androgynous; peri- 
gynia 3-3.5 mm. long, the beak subentire, 0.3-0.6 mm. long. 

C. tojquianensis 

Scales ferruginous; terminal and upper spikes androgynous, the 
others pistillate; perigynia 5-5.5 mm. long, the elongated beak 
2.5 mm. long, conspicuously bidentate at apex. 

C. tunimanensis 
Pistillate spikes mostly 6-12 cm. long; achenes in age spreading or 

conspicuously outcurved C. Donnell-Smithii 

Scales of the spikes pale, usually green or greenish, the spikes not dark- 
colored. 
Leaves 6 mm. wide or narrower. 

Leaf sheaths and bracts more or less minutely pubescent. 

C. anisostachys 
Leaf sheaths and bracts glabrous. 

Culms pale green at base; spikes erect, crowded . . .C. quichensis 
Culms purplish at base; spikes, at least the lower, drooping, 

remote C. venosivaginata 

Leaves all or chiefly 8-24 mm. wide. 

Leaves about 5-15 mm. wide; scales abruptly long-aristate. 

C. huehueteca 
Leaves 15-24 mm. wide; scales short-aristate C. Sleyermarkii 

Carex anisostachys Liebm. Dansk. Vid. Selsk. Skrivt. V. 2: 
B66. 1850. 

Mountain forest, often with Juniperus, often on open grassy 
lopes or in alpine meadows, frequently on limestone, 3300-3700 
neters; Huehuetenango (Sierra de los Cuchumatanes; collected at 
everal localities). Central and southern Mexico. 

Plants usually densely cespitose, from densely matted, woody rootstocks, the 
ulms mostly 50 cm. high or often much lower, slender but stiff, much longer than 
he leaves; basal leaf sheaths becoming shredded in age; leaves mostly 5-15 cm. 



102 FIELDIANA: BOTANY, VOLUME 24 

long, 1-3 mm. wide, flat with revolute margins, glabrous or sparsely pubescent, 
the sheaths somewhat pubescent, reddish-spotted ventrally; spikes 3-5, erect, 
close together or the lowest somewhat remote, the lowest spike conspicuously pe- 
dunculate, the upper ones short-pedunculate or subsessile, the upper spike androgy- 
nous or staminate throughout or pistillate in the center, the other spikes pistillate, 
linear, 7-15 mm. long, 3 mm. broad, the perigynia 10-20; lowest bract 2-4 cm. 
long, shorter than the inflorescence, the other bracts much shorter or absent, 
sheathless or nearly so; staminate scales hyaline with a green midrib, tinged with 
brownish or straw-color; pistillate scales ovate, narrower than the perigynia, longer 
than the perigynia and acuminate, to shorter than the perigynia and subobtuse, 
greenish-hyaline and somewhat tinged with reddish brown; perigynia rather nar- 
rowly oval-obovoid, 2.5-3 mm. long, triangular in cross section, not at all inflated, 
glabrous, with few slender nerves, green, abruptly contracted into a short beak; 
achene 1.5 mm. long, oblong-oval, acutely triangular in cross section, brownish, 
with concave sides, geniculate with the very short style; stigmas 3. 

Carex cuchumatanensis Standl. & Steyerm. Ceiba 4: 62. 
1953. 

In swampy ground along streams or in running water, Sierra de 
los Cuchumatanes, 2100-3500 meters; endemic; Huehuetenango 
(type collected in vicinity of Tunima, Sierra de los Cuchumatanes, 
3400-3500 meters, July 7, 1942, Steyermark 48347; in Herbarium of 
Chicago Natural History Museum). 

Loosely cespitose, stoloniferous and forming colonies, the stolons stout, scaly, 
and ascending, the culms 1.5-4.2 dm. high, 3-4 mm. thick at base, erect, strict, 
triangular with concave sides, smoothish or only slightly roughened above, pale 
brown at base, the lower and basal sheaths not at all filamentose; sterile shoots 
elongate, conspicuous; leaves with well-developed blades 4-5 to a fertile culm, on 
the lower third, the blades gray-green or light green, 3-4 dm. long, 3-4.5 mm. wide, 
long-tapering, closely serrulate on margins; staminate spike solitary, mostly pistil- I 
late, short-peduncled, linear-spatulate, 3.2-3.5 cm. long, 3-5 mm. wide, the scales 
oblong-obovate, obtuse, purplish-black with light-colored center not extending to 
tip and very narrow white-hyaline apex and margins; pistillate spikes 3-6, approxi- 
mate, sessile or the lower as much as 1 cm. long-pedunculate, linear, 1.8-3.2 cm. 
long, 3-5.5 mm. wide, densely flowered throughout, 100-150-flowered, appressed; 
bracts with prolonged sheaths, leafy, equaling or mostly surpassing the inflores- 
cence, greenish, pale or drab brown at base; pistillate scales ovate-oblong, the 
middle and upper ones obtuse, the lower mucronate to aristate, narrower and 
shorter than the perigynia, purplish-black with pale 1-3-nerved center 0.4-0.6 mm. 
wide not extending to tip and very narrow hyaline apex; perigynia yellowish-green 
or stramineous, broadly elliptic-ovate, plano-convex, 2.7-2.9 mm. long, 2-2.2 mm. 1 
wide, minutely serrulate at apex, membranous, papillate, round-tapering at base, I 
short-stipitate, glabrous, 6-10-nerved on both faces, abruptly apiculate, the beak 
only 0.1-0.2 mm. long, the orifice subentire or only slightly bidenticulate; achenes { 
lenticular, obovate, body of achene 1.8-1.9 mm. long, 1.5 mm. wide, the slender 
style 0.5 mm. long; stigmas 2, slender. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 103 

Carex Donnell-Smithii L. H. Bailey, Mem. Torrey Club 1: 
56. 1889. 

Usually in damp or wet, mixed, mountain forest, 1300-3800 me- 
ters; Alta Verapaz (type from Pansamald, Tuerckheim 659); Baja 
Verapaz; El Progreso; Zacapa; Jalapa; Jutiapa; Chimaltenango; 
Solola; Quich^ ; Quezaltenango; San Marcos. Honduras; El Salva- 
dor; Costa Rica; Panama. 

Plants tall and coarse, often a meter high and forming large culms; culms pur- 
plish at the base; leaves numerous, elongate, 3-10 mm. wide, glabrous, stiff, long- 
attenuate; inflorescence compound, 20-60 cm. long, the lower branches in 2's or 
4's, the upper simple; spikes numerous, androgynous, narrowly linear-cylindric, 
3-12 cm. long, 3.5-5 mm. thick, the pistillate portion loosely 20-50-flowered; scales 
lance-ovate, usually rough-awned, purplish black; perigynia lanceolate, 3.5-4 mm. 
long, membranaceous, dull green, scarcely inflated, glabrous, puncticulate, 2-costate 
and strongly few-nerved, short-stipitate, abruptly rostrate, the beak excurved, 
1-1.5 mm. long, bidentate. 

A forest plant, abundant in many localities, especially in the 
Occidente. It sometimes forms large colonies along the white-sand 
quebradas in the region between San Martin, Chile Verde and Co- 
lomba. Guatemalan material has been referred to C. Jamesonii 
Boott, a South American species. 

Carex huehueteca Standl. & Steyerm. Field Mus. Bot. 23: 195. 
1947. 

Known only from the type, Huehuetenango, Sierra de los Cuchu- 
matanes, Cerro Canana, near Canana, 2500-2800 meters, growing 
along a stream, Steyermark 49055. 

Plants densely cespitose, with short stout stolons, the culms slender, erect, 
60 cm. high or more, obtusely trigonous, dark purplish brown below the nodes, 
smooth; leaf sheaths broad and lax, dark reddish brown or purple-brown; leaves 
numerous, rather thin and soft, pale green when dried, the blades 12-27 cm. long, 
about 5-15 mm. wide, narrowly attenuate, conspicuously costate, inconspicuously 
septate between the nerves, flat, the margins smooth; spikes few or several, on very 
long, slender peduncles, apparently pendulous, the terminal one staminate, the 
others pistillate or perhaps some of them androgynous, linear, mostly 3-4.5 cm. 
long, laxly and rather distantly flowered, the bracts filiform, erect, short, long- 
vaginate at the base, the sheaths as much as 1 cm. long, purple-brown; scales pale 
brownish or brownish green or yellowish green, oblong-oval, about 2.5 mm. long, 
abruptly contracted at the apex and bearing a stiff erect awn as much as 5 mm. 
long but usually shorter; perigynia fusiform-lanceolate, greenish, com pressed- tri- 
angular in cross section, conspicuously nerved, glabrous, 3.5 mm. long, 1 mm. 
broad, gradually attenuate into the beak; stigmas 3. 

The type specimen is in poor condition for study, being in ad- 
vanced fruit, but the plant is clearly different from any other species 



104 FIELDIANA: BOTANY, VOLUME 24 

known from Central America or Mexico. It is noteworthy for its 
wide leaves, the foliage in general being reminiscent of Luzula 
gigantea. 

Carex Longii Mackenzie, Bull. Torrey Club 49: 372. 1922. 

Marshes and wet meadows, 1500-2300 meters; Alta Verapaz; 
Jalapa. United States and Mexico; Costa Rica; Venezuela. 

Plants densely cespitose, the culms 30-75 cm. tall, slender but stiff; leaves 
2-3.5 mm. wide, with green sheaths; spikes 3-10, gynaecandrous, silvery-greenish 
or silvery-brownish, aggregated into a head 2-4 cm. long, the spikes ovoid, 6-10 
mm. long and 4-6 mm. wide, obtuse; perigynia numerous, appressed-ascending; 
scales ovate, subobtuse, shorter than the perigynia; perigynia strongly compressed, 
plano-convex, 3-4 mm. long, 2 mm. wide, glabrous, the body broadly obovate, 
conspicuously winged and serrulate, abruptly short-rostrate, the beak one-third as 
long as the body, bidentate; achenes substipitate. 

This has been reported for Guatemala under the name Carex 
albolutescens Schwein. 

Carex peucophila Holm, Contr. U. S. Nat. Herb. 8: 290. pi. 
63, f. a-e. 1905. 

Moist or wet, alpine meadows or rocky slopes or cliffs, often or 
usually on limestone, 3300-3750 meters; Huehuetenango (Sierra de 
los Cuchumatanes; collected at several localities). Southern Mexico. 

Plants loosely cespitose, the rootstocks long-creeping, brownish, rather slender; 
culms slender, 40 cm. high or lower, erect or nearly so, obtusely tetragonous, much 
exceeding the leaves; leaf blades erect or ascending, 3-20 cm. long, 1.5-2.5 mm. 
wide, dull green, firm, with revolute margins; spikes 3-6, gynaecandrous, close 
together or the lowest ones separated, ovoid or subglobose, 6-9 mm. long, 4.5-6 
mm. broad, rounded at the apex, densely flowered, the staminate flowers very few 
and inconspicuous; perigynia appressed-ascending, the beaks not prominent; scales 
ovate, obtuse, somewhat narrower and shorter than the perigynia, chestnut-brown, 
with a green center; perigynia plano-convex, ovate, 4.5 mm. long, 1.8-2 mm. wide, 
narrowly wing-margined, serrulate to the middle, dull green or finally brownish, 
subcoriaceous, rounded or truncate at the base, tapering or rather abruptly con- 
tracted into a flat beak 1 mm. long; achenes lenticular, suborbicular-quadrate, 
nearly 2 mm. long, 1.5 mm. wide, yellowish, truncate at the apex, geniculate with 
the slender style; stigmas 2. 

Carex planostachys Kunze, Suppl. Riedgr. 138. pi. 35. 1840. 

Known only in Guatemala from the department of Huehuete- 
nango, where collected in dry soil near Chacula at 1600 meters, by 
Seler. Mexico to northeastern Texas. 

Plants low, from short stout rhizomes, the culms 20 cm. high or less, rough, 
brownish at the base; leaves clustered above the base, 1-2.5 mm. wide, long-atten- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 105 



uate, rough above; terminal spikes staminate, linear, 8-12 mm. long, short-pedun- 
culate, the lateral spikes pistillate, the upper 2 contiguous, sessile, the lower ones 
slender-pedunculate, basal, oblong, 5-10 mm. long, 3-4 mm. wide, 6-12-flowered; 
scales lance-ovate, acutely carinate, acute to cuspidate; perigynia obovoid-elliptic, 
3.5 mm. long, not inflated, many-nerved, light green, substipitate, abruptly ros- 
trate, the beak 0.5 mm. long, obliquely cut. 

This was reported from Guatemala by Loesener under the name 
C. Halleriana Asso. 




FIG. 24. Carex polystachya. a, Upper portion of flowering branch (X 
6, Spikelet ( X 5). c, Perigynium ( X 10). d, Scale ( X 10). e, Part of leaf ( X 

Carex polystachya Swartz ex Wahl. Kongl. Akad. Handl. 24: 
149. 1803. C. cladostachya Wahl. loc. cit. C. Hartwegii Boott ex 
Benth. PL Hartw. 96. 1848 (type collected near Guatemala, Hart- 
weg 628}. C. polystachya var. minor Boott, 111. Carex 490, 492. 1867. 
Figure 24. 



106 FIELDIANA: BOTANY, VOLUME 24 

Damp or wet, mixed forest or oak or pine forest, sometimes in 
thickets or open meadows, abundant in many localities, 300-3500 
meters, most plentiful at middle or rather high elevations; Alta Vera- 
paz; Zacapa; Chiquimula; Jalapa; Santa Rosa; Guatemala; Sacate- 
pe"quez; Chimaltenango; Solola; Quiche"; Huehuetenango; Quezalte- 
nango. Mexico to Panama, southward to Bolivia; West Indies. 

Plants erect or ascending, often forming dense clumps, with short, somewhat 
woody rhizomes, the slender culms 20-50 cm. tall; leaves numerous, clustered at 
the base of the plant, very rough, 2-7 mm. wide; spikes very numerous, in decom- 
pound panicles, androgynous, 5-10 mm. long, 3-4 mm. wide; scales ovate, striate, 
cuspidate or acute; perigynia narrowly obovoid, sharply trigonous in cross section, 
not inflated, bright green, bicarinate, many-nerved, stipitate, abruptly rostrate, 
the beak strongly bidentate, half as long as the body or shorter. 

This is the only species of Carex that is common in Central Amer- 
ica, and it is found in all the countries of Central America. 

Carex polystachya var. Bartlettii (O'Neill) Standl. & Steyerm. 
Ceiba 4: 68. 1953. C. Bartlettii O'Neill, Carnegie Inst. Wash. Publ. 
522 : 255. f. 1. 1940. 

Type from El Cayo, British Honduras, at edge of mountain pine 
ridge, Bartlett 1171 8 A; collected also near Vaca, El Cayo District, 
Gentle 2321 in part. In Guatemala known only from rocky forested 
banks, at 350 meters, Alta Verapaz (vicinity of Secanquim, Maxon 
& Hay 3214~). 

Differing from C. polystachya only in having scaberulous peri- 
gynia. 

The position of this plant still is obscure because of the lack of 
adequate material, but even so, it can scarcely be regarded as more 
than a variety of C. polystachya, and it is possible that it should be 
reduced to synonymy under C. scabrella. However, the general ap- 
pearance of the plant and its gross characters seem to ally it rather 
with C. polystachya. It is significant that Gentle 2321 consists of 
plants with glabrous perigynia and others with scaberulous ones, 
indicating, probably, that the two forms were growing together. 

Carex praegracilis Boott, Bot. Gaz. 9: 87. 1884. 

Moist meadows or in boggy borders of streams, 2100-3500 meters; 
Huehuetenango (Sierra de los Cuchumatanes: Cerro Chemalito; 
Tunima; Nuca). Western United States; Mexico; South America. 

Plants with stout long-creeping blackish rhizomes, the culms 20-70 cm. high, 
arising singly or in small clumps, acutely trigonous, slender but stiff, usually rough- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 107 

ened on the angles above, generally longer than the leaves; leaf blades erect- 
ascending, 10-25 cm. long, 1.5-3 mm. wide, light green, rather thick, rough on the 
margins; spikes 5-15, densely aggregate, all crowded together or the lowest slightly 
separated, subobtuse, 4-8 mm. long, 4-6 mm. broad, the terminal staminate 
flowers inconspicuous; scales ovate, light chestnut-brown with conspicuous hya- 
line margins and a lighter midrib, the lower ones cuspidate, the upper acuminate, 
wider and longer than the perigynia and usually completely concealing them; 
perigynia plano-convex, ovate or ovate-lanceolate, 3-4 mm. long, 1.5 mm. wide, 
erect-ascending, stramineous or at maturity brownish black, slightly several-nerved 
dorsally, sharp-edged, the margins serrulate above the middle, tapering into a ser- 
rulate beak half as long as the body of the achene or longer; achenes lenticular, 
obovoid, about 1.2 mm. long; stigmas 2. 

Carex quichensis F. J. Hermann, Journ. Wash. Acad. Sci. 40: 
284-285. 1950. 

Edge of brook, 2333 meters; endemic; El Quiche" (type collected 
above Nebaj, A. J. Sharp 45144)- 

Densely cespitose from slender, pale brown rootstocks; culms 1.8-4 dm. high, 
erect, shorter than the leaves, smooth, bluntly triangular; sterile shoots conspicu- 
ously developed; leaves 3-5 to a fertile culm, the blades flat, linear, erect, 9-20 cm. 
long, 3.5-6 mm. wide, scaberulous on the margins toward the apex, the lower 
sheaths loose, hyaline and soon breaking ventrally, the ligule not conspicuously 
prolonged, blunt, as long or twice as long as wide, 14-25 mm. long, 1.5-2 mm. wide, 
terminal spike staminate, linear, erect, 14-25 mm. long, 1.5-2 mm. wide, borne on 
a peduncle 3-5 mm. long, the scales elliptic-oblong, 4 mm. long, appressed, hya- 
line, tinged with reddish brown, the broad dark green, often somewhat carinate 
midrib smooth and extended as a mucro; lateral spikes 3-4, erect, pistillate or often 
androgynous, the lowermost widely separate on a capillary peduncle, the upper 
short-peduncled, linear, 20-30 mm. long, 3.5-5 mm. wide at maturity, 15-30- 
flowered, the perigynia appressed-ascending; bracts leaf-like, overtopping the in- 
florescence, the blade of the lowermost 15-20 cm. long, the sheaths, except the 
uppermost, very long (to 45 mm.), prolonged at the mouth and with a prominent 
ligule; scales broadly elliptic-lanceolate, acuminate, 3 mm. long, nearly equaling 
the perigynia, thin and hyaline, the margins streaked with dark red, the midrib 
stout, green, generally more or less carinate, extending to or prolonged slightly 
beyond the tip; perigynia broadly ellipsoid to narrowly obovoid, 3.2 mm. long, 
1.3-1.5 mm. wide, sessile, conspicuously many-nerved, subcoriaceous, little in- 
flated, olive-green, glabrous, tapering at the base, abruptly contracted above into 
a short (0.35 mm.), stout, straight, subentire beak conspicuously thickened at the 
mouth; achenes trigonous, narrowly obovoid, 2 mm. long, 1 mm. wide, stramineous, 
granular, tapering to the stipitate base, rounded at the apex, bent-apiculate and 
jointed with the slender style; stigmas 3, short, dark red. 

Carex scabrella Wahl. Kongl. Akad. Handl. 24: 149. 1803. 

Known only from rocky slopes, Jalapa, Potrero Carrillo north- 
east of Jalapa, 1500-1700 meters, Steyermark 33117. Greater An- 
tilles. 



108 FIELDIANA: BOTANY, VOLUME 24 

Plants densely cespitose, from thick hard rootstocks, the weak, very slender 
culms 10-40 cm. tall; leaves very numerous, clustered at the base of the plant, 
pale green, rather thin, long-attenuate, 1.5 mm. wide or less; spikelets usually 
numerous, in spike-like panicles, the upper panicles sessile, the lower long-peduncu- 
late, the spikelets androgynous, each with 3-6 ascending perigynia; scales ovate, 
acutish, often awned, several-nerved; perigynia 3-4 mm. long, obovoid, acutely 
trigonous in cross section, not inflated, conspicuously costate, tapering to the base, 
narrowed into a bidentate beak about 1 mm. long. 

The Alta Verapaz specimen (vicinity of Secanquim, Maxon & 
Hay 3214), once determined by Mackenzie as C. scabrella, has been 
examined by the writers and seems instead referable to C. poly-^ 
stachya. 

Carex Steyermarkii Standl. Field Mus. Bot. 23: 196. 1947. 

Known only from the type, Huehuetenango, Cerro Huitz, be- 
tween Barillas and Mimanhuitz, Sierra de los Cuchumatanes, 1600- 
2600 meters, Steyermark 48542. 

Plants apparently cespitose, the culms stout, erect, about 50 cm. high, striate, 
smooth; leaves very large and broad, numerous about the base of the culm and 
crowded, apparently erect, as much as 40 cm. long, about 16-24 mm. wide, flat, 
rather thick, pale green when dried, somewhat lustrous above, paler beneath, with 
a conspicuous costa, smooth on the margins, somewhat dilated and sheathing at 
the base, deep brown-red near the base; spikes rather numerous and forming a large 
lax inflorescence, pendulous on long slender peduncles, androgynous, elongate- 
linear, as much as 6 cm. long, laxly flowered or the lower flowers remote; scales 
lance-oblong, acute, inconspicuously mucronate or muticous, pale sordid brownish 
or pale greenish brown; perigynia fusiform-lanceolate, about 5 mm. long and 
slightly more than 1 mm. broad, pale sordid brown, conspicuously nerved, lus- 
trous, gradually attenuate to a short stout erect beak, obtusely triangular in cross 
section; stigmas 3. 

The type specimen consists of a single much-weathered plant, in 
which it is difficult if not impossible to determine the true characters 
of the spikes and the general form of the inflorescence. The species 
is evidently a distinct one, outstanding because of its extremely 
broad leaves, among the widest to be found in the genus. It is be- 
lieved that the relationship is with C. huehueteca Standl. & Steyerm. 

Carex Thurberi Dewey in Torr. Bot. Mex. Bound. Surv. 232. 
1859. 

In marshes, 1350-2500 meters; Alta Verapaz (near San Cristobal, 
Tactic, and Coban) ; Baja Verapaz (region of Fatal) ; Huehuetenango. 
Arizona and Mexico; Dominican Republic. 

Plants rather robust, from short thick rootstocks, the stout culms 60-100 cm. 
tall, the basal sheaths purplish and filamentose; leaves numerous, strongly septate- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 109 

nodulose, the blades often 30-60 cm. long, 8-18 mm. wide, light green, rough; 
terminal spike staminate, linear, 2-3 cm. long, the lateral spikes 3-6, approximate, 
spreading or drooping, oblong-cylindric, light green, densely many-flowered; scales 
ovate-lanceolate, rough-awned; perigynia ovoid-lanceolate, 4.5 mm. long, trigo- 
nous, scarcely inflated, subcoriaceous, strongly costate, obliquely short-stipitate, 
tapering into a strongly bidentate beak 2 mm. long, its teeth erect, stiff, 1 mm. 
long. 

Abundant about the margins of the swamps of the Fatal region. 
The spikelets, like those of C. polystachya, often are attacked by a 
smut. This species has been reported from Guatemala under the 
name C. polysticha Boeckl. 

Carex tojquianensis Standl. & Steyerm. Ceiba 4: 64. 1953. 

Dry rocky grassy slopes, Sierra de los Cuchumatanes, 3700 me- 
ters; endemic; Huehuetenango (type collected between Tojquia and 
Caxin bluff, summit of Sierra de los Cuchumatanes, 3700 meters, 
August 6, 1942, Steyermark 50150; in Herbarium of Chicago Natural 
History Museum. Paratypes: same locality, on Caxin bluff, August 
6, 1942, Steyermark 50175, 50176; in Herbarium of Chicago Natural 
History Museum). 

Loosely cespitose and substoloniferous, the stolons stout with dark blackish- 
brown or purplish-black scales, the culms 2.5-5.5 dm. high, 2-3 mm. thick below 
middle, erect, triangular, glabrous, exceeding the leaves, the lower and basal 
sheaths not filamentose; leaves with well-developed blades 7-10 to a fertile culm, 
coriaceous, stiff, strongly keeled beneath and channeled above, subrevolute, 1-2.5 
dm. long, 2.5-4 mm. wide, glabrous, long-attenuate, the sheaths dark blackish- 
brown or brown-purple at base, striate; inflorescence 5-12 cm. long, spikes 3-4, 
androgynous, usually single, spreading or drooping on unequal, elongated, slender, 
smooth peduncles mostly longer than the spikes, the spikes linear-oblong, 2-3.5 cm. 
long, 4-6 mm. wide, closely and densely flowered, the upper one-fourth or one-fifth 
staminate, the perigynia appressed-ascending; lower bract leaf-like, long-sheathing, 
7-11 cm. long, 1-1.5 mm. wide, the upper short-sheathing; pistillate scales oblong- 
lanceolate, obtuse to subacute, dark brown-purple with prominently nerved pale 
green center and very narrow hyaline margins, about the same width or slightly 
narrower than and about equaling the length or slightly exceeding the perigynia, 
3-3.25 (-5) mm. long, 1.25-1.5 mm. wide; staminate scales similar; perigynia ellip- 
tic-oblong or elliptic-lanceolate, compressed-triangular, not inflated, 3-3.5 mm. 
long, 1.25-1.5 mm. wide, stramineous or buff -greenish throughout or prominently 
reddish-brown dotted in lower half, 2-keeled, the mature perigynia strongly or 
faintly several-nerved on one or more sides, sometimes in the lower portion only, 
otherwise nerveless, narrowed at the base, substipitate, glabrous or the uppermost 
part minutely hispidulous on margins, tapering into the oblique subentire beak, 
0.3-0.6 mm. long; achenes elliptic-oblong, 2.25-2.5 mm. long, 1.25 mm. wide, 
nearly filling the perigynium body, triangular, rounded at base, apiculate, jointed 
with the slender style; stigmas 3, brownish. 



110 FIELDIANA: BOTANY, VOLUME 24 

Carex tunimanensis Standl. & Steyerm. Ceiba 4: 65. 1953. 

Open alpine meadows, along rivulets, Sierra de los Cuchuma- 
tanes, 3300-3500 meters; endemic; Huehuetenango (type collected 
in alpine areas in vicinity of Tunima, Sierra de los Cuchumatanes, 
3400-3500 meters, July 7, 1942, Steyermark 48334; in Herbarium of 
Chicago Natural History Museum. Paratypes: near Tunima, July 6, 
1942, Steyermark 48283; along rivulet, alpine meadows just north- 
west of Chancol, July 6, 1942, Steyermark 48251; in Herbarium of 
Chicago Natural History Museum). 

Cespitose, the culms 0.9-2.5 dm. high, 1-1.5 mm. thick below the middle, erect, 
triangular, glabrous, exceeding the leaves; leaves numerous, mostly clustered at the 
base of the culm, the blades stiff, the lower spreading, 5-10 cm. long, 2.5-5.5 mm. 
wide, the margins rough-serrulate, sheaths brown and conspicuously striate at 
base, spreading in age; inflorescence up to 3.5 dm. long, spikes 4-7, the terminal 
and uppermost ones androgynous with few pistillate flowers, the others pistillate, 
single or in 2's, at first erect, in maturity ascending, eventually remotely separated 
on elongated, slender, straight or flexuous, glabrous peduncles which are solitary 
or in pairs and in maturity mostly exceed the spikes, the lower peduncles at ma- 
turity eventually loosely spreading or cernuous, the spikes oblanceolate-clavate to 
clavate-cylindric, densely flowered, 1.5-3 cm. long, 4-8 mm. wide; the terminal 
and uppermost spikes erect, on peduncles 1-1.5 cm. long, the staminate scales 
membranaceous, lustrous, obovate, acute, 5-5.5 mm. long, 1.5-2 mm. wide, ap- 
pressed-ascending, ferruginous with a pale green carinate center, the margins 
white-scarious; pistillate spikes on long-exserted, slender, capillary peduncles up 
to 8 cm. long, exceeding the spikes, the perigynia ascending; bracts long-sheathing, 
with a prolonged closed and tubular sheath 15-26 mm. long and prolonged at the 
brown orifice with a conspicuous ligule, the lowest ones 6-9 cm. long, leaf-like, the 
upper ones greatly reduced; perigynia fusiform, 5-5.5 mm. long, 1.25-1.5 mm. wide, 
trigonous and 3-ribbed, slightly 1-nerved on at least two sides, coriaceous, pale 
green, glabrous, tapering into a conspicuous beak 2.5 mm. long, conspicuously 
bidentate at the apex, the hyaline teeth 0.6-0.7 mm. long; achene trigonous, ob- 
long-elliptic, 2 mm. long, 1 mm. wide, stramineous, not filling the perigynium, 
jointed with the style; stigmas 3. 

Carex venosivaginata Standl. & Steyerm. Ceiba 4: 67-68. 
1953. 

On mossy bluffs in upper reaches of barrancos and mossy ground 
in cloud forests, Sierra de los Cuchumatanes, 2400-2600 meters; en- 
demic; Huehuetenango (type collected in cloud forest, Cerro Huitz, 
between Mimanhuitz and Yulhuitz, Sierra de los Cuchumatanes, 
2600 meters, July 14, 1942, Steyermark 48554; in Herbarium of Chi- 
cago Natural History Museum. Paratype: on high bluffs in upper 
reaches of barranco, above San Juan Ixcoy, Sierra de los Cuchuma- 
tanes, 2400 meters, August 4, 1942, Steyermark 50062; in Herbarium 
of Chicago Natural History Museum). 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 111 

Loosely cespitose with short rootstocks, the culms 2.5-6 dm. high, 1-1.5 mm. 
thick below middle, slender at base, erect, triangular, the angles and sides smooth, 
strongly purple-tinged at base, the lower sheaths breaking and very slightly fila- 
mentose; leaves with well-developed blades 5-10 to a fertile culm, septate-nodulose, 
the lower clustered at the base, the blades erect, firmly membranaceous to subcoria- 
ceous, septate-nodulose, flat with subrevolute scabrous margins, 1-1.5 dm. long, 
2.5-4.5 mm. wide, the sheaths glabrous; spikes 4-6, mostly appearing androgynous, 
widely separate, in pairs or single, elongate-linear, 1.5-4 cm. long, 3.5-4 mm. wide, 
closely flowered above, loosely at base, the upper fourth to fifth staminate; lower 
bract leaf-like, long-sheathing, exceeded by the culm, the upper much reduced; 
staminate scales membranaceous, obovate, mucronate to abruptly acuminate, 4- 
4.25 mm. long, 1.5 mm. wide; pistillate scales membranaceous, broadly ovate to 
suborbicular-ovate, abruptly acuminate, 2.6-3 mm. long, about 2 mm. wide, sub- 
cucullate, carinate; perigynia narrowly oblong-elliptic, 3 mm. long, about 1 mm. 
wide, obtusely trigonous, tapering to a somewhat rounded base, glabrous, tapering 
into a smooth short beak 0.8-1 mm. long, obliquely cut, at length minutely biden- 
tate, puncticulate, 2-ribbed, strongly 3-4-nerved on each face with elevated nerves; 
achenes elliptic-oblong, 1.9-2 mm. long, 1 mm. wide, trigonous, rufous-brown, fill- 
ing about four-fifths of the perigynium body, apiculate, jointed with the slender 
style; stigmas 3. 

This species may be related to C. perlonga Fernald, from which it 
differs in the glabrous septate-nodulose sheaths, culms, and leaves 
(except for their serrulate margins), shorter perigynia with shorter 
beaks, and more remote spikes. 

Carex xalapensis Kunth, Enum. PL 2: 380. 1837. C. involu- 
cratella Mackenzie, Cariceae, N. Amer. Fl. 18: 50-51. 1931. 

At 2000 meters; Huehuetenango ("Los Pinitos," southeast of 
Huehuetenango, Steyermark 48131}. Central Mexico. 

Plants cespitose, the rootstocks short-creeping, fibrillose, the culms 1.8-1.9 dm. 
high, rather slender, 2-3 mm. thick and dull brown at base, sharply triangular, 
roughened above, equaling to exceeding the leaves; leaves with well-developed 
blades 5-8 to a culm, all on the lower third, the blades stiff, 1-3.5 dm. long, 2-5 
mm. wide, roughened on margins, the sheaths tight, not septate-nodulose dorsally 
or cross-rugulose ventrally; spikes closely aggregated into an ovoid to broadly ob- 
long head 1.5-3.5 cm. long and 0.9-1.5 cm. thick, the individual spikes 6-20, 
scarcely distinguishable, androgynous, with 10-20 appressed to spreading peri- 
gynia, the staminate flowers very inconspicuous; 1-4 of the lower bracts developed, 
setaceous, slightly enlarged at base, 1-4 cm. long, usually shorter than the inflores- 
cence; pistillate scales ovate, acuminate or short-cuspidate, somewhat narrower 
than and about length of bodies of perigynia, reddish- or yellowish-brown, with a 
3-nerved green center and narrow hyaline margins; perigynia plano-convex, the 
main body ovate to suborbicular, light green or yellowish-brown at maturity, 
3-4.25 mm. long, 2-3 mm. wide, nerveless ventrally, faintly few-nerved dorsally, 
substipitate to definitely abruptly short-stipitate at the truncate to rounded base, 
serrulate above the middle, abruptly narrowed into a serrulate, narrowly bidentate 
beak about half the length of the body; achenes lenticular, filling perigynium body, 
quadrate-suborbicular, 1.5-2 mm. long, geniculate with the slender style; stigmas 2. 



112 FIELDIANA: BOTANY, VOLUME 24 

In the single Guatemalan specimen cited, the bases of the peri- 
gynia are more markedly stipitate than in other collections of C. 
xalapensis, but there is variation in this character. Carex involucra- 
tella Mackenzie apparently represents only a more luxuriant form of 
C. xalapensis, having somewhat larger perigynia, inflorescences, and 
bracts, but is believed to represent the type of variation normally 
occurring in C. xalapensis. 

CLADIUM R. Brown 

Tall coarse perennials with thick rhizomes; leaves long and narrow, rough- 
margined; flowers perfect, small, dark brown or blackish, fasciculate along the 
branches of a terminal panicle, axillary panicles also often present; spikelets mostly 
1-3-flowered, usually only the lowest flower fertile; scales imbricate, the lowest 
usually empty; perianth represented by slender bristles, or these absent; stamens 
normally 3; style continuous with the ovary, dilated at the base, the basal portion 
sometimes persistent as a tubercle, the style branches 3; achene oblong or ovoid, 
somewhat drupe-like, smooth, ecostate or trigonous. 

About 30 species in tropical and temperate regions of both hemi- 
spheres. Only one species known from Central America. 

Cladium jamaicense Crantz, Inst. Herb. 1: 362. 1766. Schoe- 
nus Cladium Swartz, Prodr. Veg. Ind. Occ. 19. 1788. S. effusus 
Swartz, loc. cit. Mariscus jamaicensis Britton in Britt. & Brown, 
111. Fl. 1: 348. 1913. Saibal (Peten, fide Lundell). 

Swampy shores of streams or in inland marshes or swampy savan- 
nas, ascending from sea level to about 1500 meters; Pete"n; Baja 
Verapaz; Alta Verapaz; Izabal; Huehuetenango. Southeastern 
United States through Mexico to British Honduras and Panama; 
West Indies and South America, and in the Old World tropics. 

Plants very coarse and stout, arising from short thick rhizomes, the culms 1-3 
meters tall, usually 1-2.5 cm. thick below, obscurely trigonous; cauline leaves 
numerous, 60-120 cm. long, 6-20 mm. wide, thick and hard, very scabrous on the 
margins, the sheaths glabrous, often reddish or ferruginous; panicles umbelliform, 
several, decompound and lax; spikelets in clusters of 2-10 or solitary at the ends of 
the ultimate branches, narrowly ovoid, acute, terete, 4-5 mm. long, bright brown, 
the lower 6-8 scales empty; perianth none; stamens 2, the anthers 3 mm. long; 
achene 2 mm. long, ovoid, acute, brown, somewhat rugose. 

The plant often grows in shallow water of swamps. There is a 
very large stand of it in the great swamp near Patal, Baja Verapaz, 
shortly above Tactic, where it constitutes the greater part of the 
vegetation of the whole swamp. The saw-like edges of the stiff leaves 
cut the skin and flesh painfully if one is careless in walking through 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 113 

the stands of the plant. In the United States it is known by the 
name "sawgrass." In Florida the tough leaves are utilized for weav- 
ing baskets and other articles, and it seems probable that they have 
been used also by the Guatemalan Indians. The Maya name is 
reported from Yucatan as "holche." 

CYPERUS L. 

Reference: Kuekenthal, Pflanzenreich IV. 20: 1-671. 1935-36. 

Annuals or perennials, with rhizomes or with fibrous roots, the culms leafy 
near the base or leafless, not truly nodulose although falsely so in C. articulatus; 
bracts one to many at the base of the inflorescence, this umbellate or head-like; 
spikelets usually 2-many-flowered, rarely 1-flowered, the scales distichous, the 
lowest 1-2 sometimes empty, the succeeding ones containing a single perfect flower, 
the uppermost sometimes sterile or empty; stamens 1-3; perianth none; style bifid 
or trifid, not enlarged at the base, usually caducous; achene lenticular or trigonous. 

About 600 species, in almost all parts of the earth but most nu- 
merous in tropical or subtropical regions. A few species besides those 
listed here are found in southern Central America. While the sub- 
genera usually recognized in the genus are rather natural ones, and in 
theory sharply defined, as a matter of fact they are often difficult of 
recognition in dried specimens. No key that may be made for sepa- 
rating the species will be found easy for use, and probably no one 
with even the best of keys can be certain of determinations in the 
majority of cases except by comparison with authentically named 
specimens. 

KEY TO THE SUBGENERA 

Style 2-cleft; achene compressed or biconvex II. Pycreus 

Style 3-cleft; achene trigonous. 

Rachilla of the spikelet persistent, not separating from the axis to which it is 

attached, not breaking up into separate joints I. Eucyperus 

Rachilla of the spikelet deciduous from the axis and articulate with it. 

Rachilla articulate only at or near the base III. Mariscus 

Rachilla breaking up into several joints IV. Torulinium 

I. Eucyperus 

Spikelets arranged in elongate spikes. 

Spikes cylindric, like the spikelets very numerous; wings of the rachilla usually 

colored. 
Spikelets usually divergent, oblong-lanceolate, compressed; rachilla narrowly 

winged C. imbricatus 

Spikelets suberect, linear, terete or subterete; rachilla broadly winged. 

Spikelets divergent; spikes 1.5-2 cm. broad C. digitatus 

Spikelets ascending; spikes about 8 mm. broad C. semiochraceus 



114 FIELDIANA: BOTANY, VOLUME 24 

Spikes broadly ovate or turbinate, like the spikelets relatively few. 

Plants tall and coarse, usually a meter high or more; scales rather remote. 

C. prolixus 
Plants small, mostly 30-50 cm. tall; scales densely imbricate. 

Plants with repent rhizomes C. rotundus 

Plants with fibrous roots C. esculenlus 

Spikelets fasciculate or digitate upon a very short rachis, never in elongate spikes. 
Plants leafless; culms spongy and when dry appearing nodose-septate. 

C. articulatus 
Plants usually leafy; culms neither spongy nor septate. 

Spikelets collected in a single dense head. 

Scales muticous ,. C. tenerrimus 

Scales mucronate. 

Plants annual; culms not thickened at the base C. humilis 

Plants perennial; culms thickened at the base. 

Scales not closely appressed, with somewhat outcurved tips. 

C. uncinulatus 
Scales appressed, the tips not outcurved. 

Spikelets white or pale yellowish; leaf sheaths blackish. 

C. seslerioides 
Spikelets golden brown; leaf sheaths purplish brown or paler. 

C. Arsenii 
Spikelets variously arranged but not in a single head. 

Plants annual. 

Spikelets green , . . . C. compressus 

Spikelets brown or castaneous. 

Culms very slender, about 1 mm. thick C. amabilis 

Culms stouter, 2-3 mm. thick C. haspan 

Plants perennial. 

Culms subterete, leafless. 

Flowers perfect; cultivated plants C. alternifolius 

Flowers dioecious; native plants C. canus 

Culms conspicuously trigonous or triquetrous. 
Scales incurved or loosely tortuous at the apex. 

Culms much shorter than the umbels C. simplex 

Culms much longer than the umbels C. Lundellii 

Scales straight, not incurved at the apex, acute or cuspidate-mucronate. 
Spikelets turgid; scales with numerous conspicuous nerves. 

C. diffusus 

Spikelets strongly compressed; scales not strongly nerved. 
Culms and leaves evidently viscid, at least in the fresh state. 

Spikelets 10-20 mm. long C. oxylepis 

Spikelets 5-8 mm. long C. elegans 

Culms and leaves not at all viscid. 

Spikelets 1-1.5 mm. wide C. haspan 

Spikelets 2 mm. wide or often much wider. 
Scales conspicuously red-lineolate, not evidently cellulose- 
reticulate; stamens 3 C. Lundellii 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 115 

Scales conspicuously cellulose-reticulate, not red-lineolate ; 

stamen 1. 

Spikelets at maturity oblong-linear to oblong-lanceolate, 
densely aggregate and radiating, never forming glom- 
erate-lobate heads. 

Leaves 2-4 mm. wide, scarcely or not septate-nodulose; 
scales rugulose; spikelets 2 mm. wide . . .C. ochraceus 
Leaves 6-10 mm. wide, conspicuously septate-nodulose; 
scales not rugulose; spikelets 2.5-3 mm. wide. 

C. virens 

Spikelets ovate or lance-ovate, densely glomerate and form- 
ing lobate heads. 

Scales oblong-lanceolate, the apex excurrent into a mi- 
nute ex curved mucro C. pseudovegetus 

Scales acute or short-mucronate, the apex straight or in- 
curved. 
Scales straight, not incurved at the apex; leaves 2-3 

mm. wide C. surinamensis 

Scales incurved at the apex; leaves 3-6 mm. wide. 

C. Luzulae 

II. Pycreus 

Spikelets 3-5 mm. wide; achene black C. unioloides 

Spikelets 1.5-2.5 mm. wide. 

Superficial cells of the achene rectangular, oblong, the achene transverse- 
undulate. 

Scales castaneous or deep brownish C. piceus 

Scales yellowish C. flavescens 

Superficial cells of the achene hexagonal, the achene (as seen beneath a lens) 

reticulate or puncticulate. 
Scales castaneous or blackish. 

Spikelets loosely spicate, the spikelets usually distant in the spikes. 

C. rivularis 

Spikelets densely spicate, the whole inflorescence head-like C. niger 

Scales greenish or yellowish, pale. 

Rachilla not winged; achene obovate, 0.6 mm. wide, rounded at the apex; 

spikelets about 2 mm. wide C. lanceolatus 

Rachilla with hyaline wings forming a collar at the base of each achene; 
achene oblong-obovate, 0.4 mm. wide, subtruncate at the apex; spike- 
lets about 1.5 mm. wide C. polyatachyos 

III. Mariscus 

Culms papillose and somewhat septate when dried; leaves very coriaceous, pale 
green; plants coarse and stout, often a meter high C. ligularis 

Culms neither papillose nor septate; leaves not thick-coriaceous. 
Plants annual, often low and 15 cm. tall or less. 

Plants mostly 15 cm. tall or less; spikelets capitate C. aristatus 

Plants usually much taller; spikelets spicate C. panamensis 



116 FIELDIANA: BOTANY, VOLUME 24 

Plants perennial, usually taller. 

Scales not closely appressed, or at least the apex not appressed. 

Spikelets oblong or lance-oblong C. Fendlerianus 

Spikelets ovate. 

Scales dark red C. Bernoullii 

Scales whitish or fulvous. 

Scales mucronulate from an excised apex and appearing tridentate, 

dirty white; culms slender, smooth C. ischnos 

Scales rounded at the apex, yellowish; culms rigid, often very rough. 

C, manimae 
Scales closely appressed for their whole length. 

Achene linear, almost 2 mm. long and 0.4 mm. wide; leaves 2-3 mm. wide; 
spikelets 4-8-flowered, only 1 mm. wide; plants usually low and 30 cm. 
tall or less C. tennis 

Achene oblong to obovoid, 0.6-1 mm. wide; leaves 3-10 mm. wide; spike- 
lets 1-30-flowered, 1-2.5 mm. wide. 

Spikes subglobose, about as broad as long C. globulosus 

Spikes cylindric, often greatly elongate, at least longer than broad. 
Scales 4-4.5 mm. long; spikelets 4-8-flowered, usually very distant on 

the rachis; leaves 2-10 mm. wide C. lentiginosus 

Scales 2-3.5 mm. long. 
Spikelets 1-3-flowered. 

Spikelets 5-10 mm. long C. hermaphroditu 

Spikelets 3-5 mm. long. 

Spikelets becoming turgid at maturity, ascending, at least in age; 
achene obovate or ovate C. flavus 

Spikelets not turgid at maturity, divaricate from the first; 
achene oblong or ellipsoid. 

Spikelets linear-lanceolate, 2.5-3 mm. long; achene oblong. 

C. pollens 

Spikelets oblong-elliptic, 3-5 mm. long; achene broadly oblong 
or ellipsoid C. Mutisii 

Spikelets containing 5 or more flowers C. hermaphroditus 

IV. Torulinium 

Achene linear; scales usually distant, the apex of one scale not reaching the base 
of the one next above on the same side of the rachilla; wings of the rachilla 
linear, reaching the top of the achene C. acicularis 

Achene oblong or obovoid; scales imbricate on the same side of the rachilla; wings 
of the rachilla elliptic, often not extending beyond the middle of the achene. 

Inflorescence consisting of a single large subglobose head C. macrocephalus 

Inflorescence with some stalked heads of spikelets. 

Spikelets divaricate, scattered along the rachis of the spike C. odoratus 

Spikelets erect or suberect, densely crowded C. Eggersii 

Cyperus acicularis (Schrad.) Steud. Syn. 45. 1855. Dididium 
aciculare Schrad. ex Nees in Mart. Fl. Bras. 2, pt. 1: 55. 1842. 
C.ferax var. acicularis Kuekenthal, Pflanzenreich IV. 20: 619. 1936. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 117 

Jutiapa, in ditch near Jutiapa, 900 meters. Yucatan and British 
Honduras; Brazil. 

Plants perennial, with a short thick rhizome, the culms 30-90 cm. tall, 3-5 mm. 
thick at the apex, stout, smooth, leafy; leaves 3-5 to each culm, 15-80 cm. long, 
5-20 mm. wide, flat, rough-margined; bracts 6-8 and 10-60 cm. long, 5-20 mm. 
wide; rays of the umbel 4-12, branched, unequal, the peduncles short or elongate; 
spikes 2-5 cm. long, 2-3.5 cm. wide, usually lax; spikelets mostly distant, divergent 
or reflexed, 5-25 mm. long, linear or acicular, the rachilla breaking up into 1-fruited 
joints, winged; scales 2 mm. long, ovate, imbricate or distant, mucronulate, brown 
or reddish brown; achene linear or linear-oblong, 1.7 mm. long, 0.3-0.4 mm. wide, 
brown, trigonous, minutely puncticulate. 

Perhaps only a form of C. odoratus (C. ferax), with which it was 
united by Kuekenthal. 

Cyperus alternifolius L. Mant. PI. 2: 28. 1771. 

Frequently cultivated for ornament, principally in fountains of 
patios. Native of Africa, but cultivated for ornament in many other 
parts of the earth, even in temperate regions. 

Plants coarse, perennial, with very short, thick rhizomes, the culms 30-150 cm. 
tall, 1-5 mm. thick at the apex, trigonous but the angles obtuse or rounded; leaves 
reduced to sheaths, these reddish brown, coarsely nerved; bracts usually 18-20 and 
10-30 cm. long, subequal, 15 mm. wide or less, rough on the margins and costa, 
deflexed at maturity; rays as many as the bracts, 2-10 cm. long, subequal, with a 
few short raylets; spikelets 3-12 in the head, ovate-lanceolate to linear-lanceolate, 
5-10 mm. long, 12-30-flowered, acute, compressed, the rachis not winged; scales 
ovate to lanceolate, imbricate, thin-chartaceous, 3-5-nerved, subacute, pale or with 
reddish brown spots; achene 0.6-0.9 mm. long, trigonous, dark brown, minutely 
cellular, short-apiculate. 

Called "tule de jardin" in El Salvador. O'Neill states that the 
species is found in Yucatan in "waste places; introduced," but there 
is no apparent evidence that it grows in Mexico and Central America 
outside gardens. It very closely resembles C. canus, and it is not 
easy to decide which species is represented unless the source of the 
specimens is known. Also in cultivation in Guatemala, apparently, 
is Cyperus Papyrus L., the giant papyrus of the Nile, from which the 
ancient Egyptians prepared a kind of paper for their manuscripts. 
That species, however, is not closely similar to C. alternifolius, but 
does resemble and is related to the native Cyperus giganteus. 

Cyperus amabilis Vahl, Enum. PL 2: 318. 1806. C. aureus 
HBK. Nov. Gen. & Sp. 1: 205. 1815. C. glareosus Liebm. Vid. Selsk. 
Skrivt. V. 2: 204. 1851. C. aureus a macrostachyus Boeckl. Linnaea 
35: 494. 1868. C. amabilis var. macrostachyus Kuekenth. Pflanzen- 
reich IV. 20: 266. 1936. 



118 FIELDI ANA: BOTANY, VOLUME 24 

Open, grassy and rocky slopes, or in wet sand along streams, 
2500 meters or less; Zacapa; Santa Rosa; Escuintla; Guatemala; 
Sacatepe"quez; Chimaltenango; Suchitepe'quez; Quezaltenango; Hue- 
huetenango. Arizona and Mexico to Costa Rica; West Indies and 
South America; tropical Asia and Africa. 

Plants annual, slender, the culms more or less cespitose, few or numerous, 
5-15 cm. tall, obscurely trigonous, smooth; leaves 1-2 near the base of the culm, 
very narrow, the sheaths reddish; bracts 3-6, foliaceous, usually shorter than the 
inflorescence; rays of the umbel 10 or fewer, simple or compound, 1-8 cm. long; 
spikelets digitately clustered at the ends of the rays, linear, 8-18 mm. long, scarcely 
more than 1 mm. wide, compressed, subacute, many-flowered, the rachilla not 
winged; scales rather laxly imbricate, spatulate, orange or ferruginous, truncate 
and excised at the apex, short-mucronulate; achene obovate or obovate-oblong, 
trigonous, brown, whitish-puncticulate, rounded at the apex and minutely apiculate. 

The Guatemalan material belongs to var. macrostachyus (Boeckl.) 
Kuekenth., in which the costa of the glumes is excurrent as a straight 
or excurved mucro. 

Cyperus arista tus Rottb. Descr. & Icon. 4: 23. 1773. C. unci- 
natus R. Br. Prodr. 215. 1810, not Poir. C. inflexus Muhl. Descr. 
Gram. 16. 1817. 

Sandy plains along river; Huehuetenango (near Cuilco). United 
States and Mexico; West Indies; South America; Old World tropics. 

Plants annual, glabrous, aromatic when drying, the culms densely cespitose or 
separated, mostly 5-15 cm. tall, about 1 mm. thick, smooth; leaves 2-6 to each 
culm, 10-15 cm. long, 2 mm. wide or less, flat, smooth on the edges, the sheaths 
purplish brown; bracts 2-5, unequal, 1.5-8 cm. long; rays few and simple, or the 
inflorescence consisting of a single head, this 5-20 mm. broad, usually very dense; 
spikelets 5-40, mostly 2-3 mm. wide, compressed, 5-16-flowered, the rachilla zig- 
zag, very narrowly winged; scales oblong-spatulate, reddish brown, 7-9-nerved, 
with hyaline margins, with a conspicuous excurved mucro; achene about 1 mm. 
long, oblong, trigonous, dark brownish purple, minutely puncticulate, stipitate, 
scarcely apiculate. 

From Yucatan there are reported the Maya names "chabxan," 
"chabxaan," and "cabaxan." 

Cyperus Arsenii O'Neill & Benedict, Leafl. West. Bot. 4: 33. 
1944. 

Dry plains or hillsides, 200 meters; Zacapa (near Estanzuela, 
Steyermark 29095). Central Mexico. 

Plants perennial, with short repent rhizomes; culms very slender, 8-15 cm. 
high, 2-3 mm. thick, minutely scaberulous at the apex, somewhat thickened at the 
base; leaves 5-10 cm. long, 0.3-0.8 mm. wide, antrorsely scabrous at the apex; 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 119 

sheaths purplish, separating in age into fibers; bracts 2, very unequal, 1-5 cm. long; 
; spikelets 5-7, forming a head 5-7 mm. in diameter, 4-5 mm. long, 1.5-2 mm. wide, 
| oblong, obtuse, subcompressed, 8-17-flowered, the rachilla not winged; scales un- 
i equal, 1.3-1.6 mm. long, acute, broadly ovate, membranaceous, scarcely mucronu- 
i late, golden brown; stamen 1; style 3-fid; achene 0.6-1 mm. long, purplish fuscous, 

trigonous, obovoid or ovoid, minutely puncticulate. 

Cyperus articulatus L. Sp. PI. 44. 1753. Canutillo; Polol 
(Pete"n, Maya). 

Swamps or marshes, most often in shallow water at the margins 
of streams or lakes, at 1500 meters or less, usually at or little above 
sea level; Pete"n; Izabal; Chiquimula; Jutiapa; Guatemala; Huehue- 
tenango. Southern United States to Mexico, British Honduras, and 
Panama; West Indies; South America; Old World tropics. 

Plants coarse and stout, with long-creeping thick rhizomes, the culms mostly 
1-2.5 meters tall, 4-15 mm. thick at the base, 2-3 mm. thick at the apex, terete, 
smooth, septate-nodose; leaves reduced to a few sheaths; bracts 2-3, scale-like, 
cuspidate, 5-15 mm. long; rays of the umbel 4-12, often branched, 12 cm. long or 
less; spikelets in clusters of 2-20, digitate, linear, 6-50 mm. long, 1 mm. wide, 
compressed, 12-40-flowered; scales brown, elliptic-ovate, obtuse, membranous, im- 
bricate, 3-7-nerved, the keel green, the wings of the rachilla lanceolate; achene 1.3- 
1.5 mm. long, 0.5 mm. broad, trigonous, oblong, dark purplish brown, apiculate. 

Called "sontul" in El Salvador. The thick spongy culms doubt- 
less are used in Guatemala for making mats and other articles. 

Cyperus Bernoullii Kuekenth. Pflanzenreich IV. 20: 464. 1936. 
Mariscus guatemalensis Clarke, Kew Bull. Add. Ser. 8: 16. 1908, not 
C. guatemalensis Steud. 1855. Type from Quezaltenango, Bernoulli 
& Carlo 1001. 

Plants perennial, with ligneous stolons 4-5 cm. long, the culms 30-40 cm. tall, 
rigid, setulose-scaberulous; leaves shorter than the culms, 3 mm. wide, very sca- 
brous; bracts 4, very scabrous, the lower ones longer than the inflorescence; inflo- 
rescence head-like, the spikes 3-4, ovate-globose, 1.5 cm. broad, very dense; 
spikelets numerous, spreading, linear-oblong, 6 mm. long, 1.7 mm. wide, subcom- 
pressed, about 10-flowered, the rachilla narrowly winged; scales rather remote, 
ovate, rounded-obtuse at the apex, dark red, faintly several-nerved; achene four- 
fifths as long as the glume, subobovate-oblong, trigonous, apiculate. 

We have seen no material of this species, which is known only 
from the original collection. 

Cyperus canus Presl, Reliq. Haenk. 1: 179. 1830. Tule; Jun- 
quillo; Sivate; Say (Coban, Kekchi). 

Damp thickets, wet fields, or frequently along stream banks, espe- 
cially among rocks, sometimes planted, 200-1500 meters; Alta Vera- 



120 FIELDIANA: BOTANY, VOLUME 24 

paz; Baja Verapaz; Zacapa; Santa Rosa; Escuintla; Guatemala; 
Chimaltenango; Quiche"; Retalhuleu; doubtless also in other depart- 
ments. Mexico to Costa Rica; Colombia. 

A glabrous perennial with short thick rhizomes, the culms robust, mostly 1-1.5 
meters tall, 4 mm. thick at the apex, 6-10 mm. thick at the base, obtusely trigo- 
nous, multistriate, smooth or very minutely scaberulous, stiff; leaves reduced to 
sheaths at the base of the culm, these purplish brown; bracts 20-30 cm. long, 
6-12 mm. wide, subequal; rays of the umbel 10-18, compound; staminate spike- 
lets in digitate clusters of 3-8 at the ends of the raylets, 6-10 mm. long, 2.5-3 mm. 
wide, 10-40-flowered; pistillate spikelets 5-14 mm. long, 2-3 mm. wide; rachilla 
wingless; scales ovate, thin, acute, closely appressed, reddish brown; achene 0.5 
mm. long and half as wide, trigonous-ellipsoid, dark brown. 

Called "tule" and "till 1 de petate" in El Salvador. The plant is 
rather scarce in the wild state, but it is frequently planted in small 
patches or fields, especially in the Pacific boca costa and about Coban. 
In Guatemala, as well as in other parts of Central America, the plant 
is of considerable utility and importance, chiefly for the weaving of 
the coarse mats used as beds. In Guatemala the mats called petates 
ordinarios are woven from it. The plant is a rather handsome and 
ornamental one, closely resembling C. alternifolius, as already re- 
marked under that species. 

Cyperus compressus L. Sp. PI. 46. 1753. 

Moist open slopes, damp thickets, or in wet sand near streams, 
at 900 meters or less; Pete"n Izabal; Zacapa; Chiquimula; Jutiapa; 
Escuintla; Suchitepe"quez; Solola. United States to Mexico, British 
Honduras, and Panama; West Indies; South America; Old World 
tropics. 

A low glabrous annual with fibrous roots, the culms mostly 5-15 cm. tall, about 
1 mm. thick, smooth; leaves 1-4 at the base of the culm, about equaling the culm, 
1.5-3 mm. wide, sometimes minutely scaberulous on the margins, the sheaths pur- 
plish brown; bracts 3-5, unequal, 1-3 mm. wide; rays of the umbel simple, 1-12 cm. 
long; spikelets in digitate clusters of 3-10, linear-oblong, 10-24 mm. long, 2-3 mm. 
wide, acute, compressed, the rachilla winged; scales firm, closely imbricate, ovate, 
3-3.5 mm. long, acuminate and mucronate, the tips somewhat excurved, 9-13- 
nerved, green, the margins broadly white-hyaline; achene obovoid, trigonous, 1-1.3 
mm. long, dark brown and shining, minutely puncticulate, subemarginate at the 
apex. 

Cyperus diffusus Vahl, Enum. PI. 2: 321. 1806. C. tolucensis 
HBK. Nov. Gen. & Sp. 1: 206. 1815. C. umbrosus Lindl. & Nees in 
Mart. Fl. Bras. 2, pt. 1: 31. 1842. C. diffusus var. umbrosus Kue- 
kenth. Pflanzenreich IV. 20: 210. 1936. C. diffusus var. tolucensis 
Kuekenth. op. cit. 211. 1936. Junquillo. 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 121 

Wet thickets or shaded banks, often in moist forest, rarely in 
open places, 1300 meters or less, most common at low elevations; 
Alta Verapaz; Izabal; Santa Rosa; Escuintla; Suchitepe'quez; Re- 
talhuleu; Quezaltenango; San Marcos. Mexico to British Honduras 
and Panama; West Indies; South America; tropical Asia and Ma- 
laysia. 

Plants perennial, glabrous, with short slender rhizomes, the slender or stout 
culms 20-50 cm. tall, about 1.5 mm. thick, trigonous, smooth; leaves 2-7 to each 
culm, shorter than or about equaling the culm, 3-9 mm. wide, flat, rough-margined, 
the sheaths reddish brown; bracts 5-8, unequal, 1-9 mm. wide; rays of the umbel 
8-12, unequal, branched; spikelets solitary or in heads of 2-3, ovate to oblong, 
4-10 mm. long, 1.5-2 mm. wide, green or pale green, 5-10-flowered, the rachilla 
not winged; scales ovate, membranous, 2-3 mm. long, cuspidate-mucronate, 7-9- 
nerved; achene 1.4 mm. long, trigonous-obovoid, dark brown, smooth, substipitate. 

A common weedy plant in lowland thickets of Guatemala, as well 
as in Central America generally. It is one of the commonest Cyperus 
species of Central America. Of the varieties and subspecies proposed 
by Kuekenthal the Guatemalan material, according to O'Neill, is 
referable to var. tolucensis (HBK.) Kuekenth. 

Cyperus digitatus Roxb. Hort. Beng. 81. 1814. 

Lake margins, Lago de Atescatempa, Jutiapa, 500 meters, Steyer- 
mark 31875. Southern Mexico; South America; Old World tropics. 

A glabrous perennial with very short rhizomes, the culms more or less cespitose, 
stout, 50-150 cm. tall, trigonous, smooth, leafy at the base; leaves about equaling 
the culms, 5-15 mm. wide, coriaceous, the sheaths purple-brown; bracts 5-7, longer 
than the inflorescence; umbels compound or decompound, with 6-10 rays; spikes 
of spikelets cylindric, 3-6 cm. long, 1.5-2 cm. broad; spikelets numerous, divari- 
cate, linear, acute, about 10 mm. long and 1-1.5 mm. wide, subcompressed-angular, 
the rachilla winged; scales densely imbricate, ovate or elliptic, membranaceous, 
stramineous, yellow, or rufous, 3-5-nerved, mucronate-cuspidate; achene one-half 
to three-fifths as long as the glume, oblong-elliptic, trigonous, fuscous, densely 
puncticulate, conspicuously apiculate. 

Cyperus Eggersii Boeckel. Gyp. Nov. 1: 53. 1888. 

Wet or marshy soil, often along streams; Izabal; Pete"n. Texas 
to Campeche and British Honduras; Honduras; West Indies; 
Ecuador. 

A coarse perennial, glabrous, with a very short, stout rhizome, the culms 20- 
90 cm. tall, 2-3 mm. thick at the apex, obtusely trigonous, sometimes rough on the 
angles; leaves 4-6 to a culm, 5-15 mm. wide, 12-60 cm. long, subcoriaceous, rough- 
margined, the sheaths reddish brown; bracts 5-8 and 4-10 mm. wide; umbel with 
5-10 rays, these unequal, usually short, with 3-5 ovate or ovate-cylindric spikes 
congested into a dense head; spikelets numerous, oblong or linear-lanceolate, 6-10 



122 FIELDIANA: BOTANY, VOLUME 24 

mm. long, about 1 mm. wide, suberect, subterete, 3-9-flowered, the rachilla break- 
ing up into 1-flowered joints; scales 2 mm. long, ovate, obtuse, sometimes mucron- 
ulate, clasping the achene, stramineous or lustrous brown; achene oblong, 1.2 mm. 
long and half as wide, obtusely trigonous, very shortly stipitate, minutely punctic- 
ulate, apiculate, yellowish brown. 

This is closely related to C. macrocephalus, from which it may not 
be more than varietally distinct. 

Cyperus elegans L. Sp. PI. ed. 2. 68. 1762. 

Retalhuleu (in marsh, Champerico, at sea level); reported from 
Escuintla and Suchitepe"quez, but perhaps incorrectly. Southern 
United States to Mexico, British Honduras, and Costa Rica; West 
Indies. 

A glabrous perennial with very short rhizomes, the culms cespitose, 30-60 cm. 
tall, 1.5-2 mm. thick at the apex, sometimes roughened and septate-nodulose, 
somewhat trigonous, the whole plant more or less viscid; leaves about as long as 
the culms, 1-4 mm. wide, involute, septate-nodulose, coriaceous, scabrous on the 
margins, the sheaths brownish; bracts 3-5, very unequal, 5-40 cm. long, 1-4 mm. 
wide; rays of the umbel 3-8, very unequal, 12 cm. long or less, sometimes com- 
pound; spikelets capitate or flabellate-capitate, 3-20 in a cluster, oblong to linear, 
3-15 mm. long, 2.5-3.5 mm. wide, 6-20-flowered, turgid; rachilla not winged; scales 
2-3 mm. long, broadly ovate to rounded, imbricate, viscid, cuspidate-mucronate, 
7-9-nerved, mostly reddish purple when young but fading to stramineous, the 
margins clasping the achene; achene trigonous-obovoid, 1.5 mm. long, evidently 
stipitate, apiculate, black and lustrous. 

Cyperus esculentus L. Sp. PI. 45. 1753. 

Zacapa, wet field near Zacapa, 200 meters; probably in other 
parts of Guatemala, but not collected. Southern Canada to Mexico 
and Panama; West Indies and South America; Old World tropics. 

A glabrous perennial with long and very slender stolons, these terminating in 
globose or ovoid tubers; culms trigonous, 15-50 cm. tall, smooth; leaves numerous, 
about equaling the culms, 3-10 mm. wide, flat, smooth on the margins, the sheaths 
pale or reddish brown; bracts 2-6, longer or shorter than the inflorescence; rays 
of the umbel 5-10, simple or compound; spikelets distant in the spike, 5-24 in each 
spike, divaricate, linear, 6-30 mm. long, 1-3 mm. wide, when mature turgid and 
not compressed, 8-40-flowered; rachilla winged; scales 2.5-4 mm. long, pale brown, 
thin, 7-9-nerved, imbricate, slightly spreading at the apex, mucronulate; achene 
1.5-2 mm. long, broadly oblong, trigonous, obtuse, not apiculate, light brown or 
grayish, lustrous, puncticulate. 

The English name is "chufa," the plant being sometimes culti- 
vated for its hard but rather sweet and agreeably flavored tubers, 
which are eaten raw. It is probably the tubers of this species which 
are offered for sale in the markets of Quezaltenango, Totonicapan, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 123 

and other places, under the name "sintule." The plant has not been 
noted in cultivation in Guatemala and the tubers are not common 
in the markets. 

Cyperus Fendlerianus Boeckel. Linnaea 35: 520. 1868. C. 
Schweinitzii var. debilis Britton, Bull. Torrey Club 13: 208. 1886. 
C. Fendlerianus var. debilis Kuekenthal, Pflanzenreich IV. 20: 465. 
1936. 

Reported by Kuekenthal from Dept. Huehuetenango, Seler 2710. 
Southwestern United States and Mexico. 

A glabrous perennial with short rhizomes, the slender culms 20-30 cm. tall, 
scaberulous above on the angles, bulbous-thickened at the base; leaves 3-4 to a 
culm, shorter than the culm or longer, 2-4 mm. wide, rough-margined; bracts 4-5, 
longer than the inflorescence, this contracted, 4-6-radiate, the rays usually short 
but rarely 6 cm. long, sometimes compound; spikes oblong-ovate, about 1 cm. long, 
containing numerous spikelets; spikelets suberect, oblong-lanceolate, acute, 4-7 
mm. long, 2 mm. wide, subcompressed, 6-8-flowered, the rachilla not winged; 
scales laxly imbricate, finally patulous at the apex, persistent, articulate at the 
base, broadly ovate, grayish and purplish-striate or dirty stramineous, obscurely 
9-11-nerved, bearing a short excurved mucro; achene four-fifths as long as the 
glume, triquetrous-ovoid, apiculate, densely punctulate. 

The Guatemalan collection is referred by Kuekenthal to var. 
debilis. 

Cyperus flavescens L. Sp. PI. 46. 1753. 

Usually in sand of stream beds, sometimes in moist thickets, at 
900 meters or less; Alta Verapaz; Zacapa; Jutiapa; Santa Rosa; Es- 
cuintla; Retalhuleu; probably in all the Pacific coast departments. 
United States to Mexico and Panama; West Indies; South America; 
Europe and Africa. 

A glabrous annual with fibrous roots, the culms slender, cespitose, mostly 5-15 
cm. tall, smooth; leaves few, shorter than the culms, about 1 mm. wide, flat, the 
sheaths purplish; bracts about 3, longer than the inflorescence; rays of the umbel 
few, simple, often very short or obsolete; spikelets 3-10 in each cluster, linear- 
oblong, subacute, 6-10 mm. long or even longer, 2 mm. wide, compressed, mostly 
10-18-flowered, the rachilla not winged; scales densely imbricate, ovate, obtuse, 
yellowish or finally brownish, often mucronulate, 3-nerved; achene one-third to 
one-half as long as the glume, broadly obovate, biconvex, brown or fuscous-brown, 
somewhat lustrous, transversely zonate, contracted at the base, apiculate. 

Cyperus flavus (Vahl) Nees, Linnaea 19: 698. 1847. Mariscus 
flavus Vahl, Enum. PI. 2: 374. 1806. Kyllinga cayennensis Lam. 111. 
1: 149. 1791. C. cayennensis auct. C. flavomariscus var. peduncu- 
laris Britton, Bull. Torrey Club 13: 215. 1886. C. flavus var. pe- 



124 FIELDIANA: BOTANY, VOLUME 24 

duncularis Kuekenth. Pflanzenreich IV. 20: 532. 1936. Sintule; 
Sacabasto. 

Moist or wet fields or thickets, sometimes a weed in cultivated 
ground, often in sand along streams, occasional in pine forest, grow- 
ing on Volcan de Pacaya among rocks of the active cone and ascend- 
ing there to 2400 meters, probably introduced by cattle, but in other 
parts of the country ascending to 2700 meters, and growing mostly 
at 1500 meters or more; Pete"n; Alta Verapaz; Zacapa; Jalapa; Jutiapa; 
Guatemala; reported from Sacatepe"quez; Chimaltenango; Huehue- 
tenango; San Marcos. Southern United States (where introduced) 
to Mexico, British Honduras, and Panama; West Indies; South 
America. 

A glabrous perennial with short thick rhizomes, the culms rather stout, 30- 
100 cm. tall, 2-3 mm. thick at the apex, trigonous, sometimes roughened on the 
angles; leaves about 8 to a culm, shorter or longer than the culm, 2-6 mm. wide, 
thin and flat, scaberulous on the margins, the sheaths reddish purple; bracts 4-6, 
unequal, mostly longer than the inflorescence; rays of the umbel sometimes 5 cm. 
long but usually much shorter and often none, simple; spikes 1-2 cm. long, 5-10 
mm. wide, oblong to ovate, very dense, with numerous spikelets; spikelets 1-2- 
flowered, 3-5 mm. long, 1 mm. wide, at least the upper ones ascending, elliptic, 
somewhat quadrangular, the rachilla winged; scales 2.5-3 mm. long, ovate-elliptic, 
9-nerved, stramineous to brown, with hyaline margins; achene about 2 mm. long, 
trigonous-ellipsoid, dark brown, minutely puncticulate. 

Called "tule" in Oaxaca. In recent years this species has been 
treated under the name Cyperus cayennensis (Lam.) Britton, but this 
name was discarded by Kuekenthal because he considered the applica- 
tion of the name Kyllinga cayennensis uncertain. It would seem that 
the proper name for the species should be C. incompletus (Jacq.) Link, 
based upon Kyllinga incompleta Jacq., published in 1790, but this 
likewise he rejected. The leaves and culms of this and other species 
of Cyperus are used commonly in Guatemala for stuffing saddle pads 
used on pack animals. 

Cyperus globulosus Aubl. PL Guian. 47. 1775. C. echinatus 
Wood, Class-book 734. 1863. 

Grassy slopes or in sand along streams, at 600 meters or less; 
reported from Pete"n; Zacapa; Chiquimula. Southeastern United 
States; Yucatan; Greater Antilles; northern South America. 

A glabrous perennial with short rhizomes, the culms slender, more or less cespi- 
tose, 15-30 cm. tall, smooth, enlarged at the base; leaves mostly shorter than the 
culms, 1.5-4 mm. wide, flat, the sheaths purplish; bracts 5-8, much exceeding the 
inflorescence; rays of the umbel 6-12, simple, 5 cm. long or shorter, the spikes sub- 
globose, 5-8 mm. in diameter, rather dense; spikelets numerous, divergent or the 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 125 

lower ones reflexed, linear-lanceolate, acute, 4-6 mm. long, 1 mm. wide, 3-6-flow- 
ered, the rachilla winged; scales rather remote, oblong-elliptic, appressed, rufous 
or stramineous, 9-nerved; achene three-fifths as long as the glume, trigonous- 
oblong, apiculate. 

This species is not reported from the Yucatan Peninsula by 
O'Neill, although collected at Sisal as long ago as 1865, and so re- 
ported by Kuekenthal. 

Cyperus guatemalensis Steud. Syn. Cyper. 47. 1855. 

Type said to have been collected in Guatemala, Kegel 1270, and 
reported also from Martinique. The collector is unknown to the 
writers in connection with Guatemala. Kuekenthal lists the species 
as doubtful or not determinable. 

Cyperus haspan L. Sp. PI. 45. 1753. C. juncoides Lam. 111. 1: 
147. 1791. C. haspan subsp. juncoides Kuekenth. Repert. Sp. Nov. 
23: 184. 1926. 

Wet savannas, marshes, or bogs, sometimes along streams or in 
ditches, ascending from sea level to 1500 meters; Alta Verapaz; 
Izabal; Chiquimula; Jalapa; Jutiapa; Santa Rosa. Southern United 
States to Mexico, British Honduras, and Panama; West Indies; 
South America; Old World tropics. 

Plants glabrous, annual or perennial, with short rhizomes or only fibrous roots, 
the culms subcespitose, 20-60 cm. tall, 2-3 mm. wide at the apex, compressed- 
trigonous to 3-winged, rather soft and weak; leaves reduced to a few purplish 
brown sheaths at the base of the culm, very rarely with blades; bracts 2, unequal, 
1-5 cm. long; rays of the umbel very unequal, usually compound, the spikelets 
clustered at the ends of the raylets, linear, acute, 4-10 mm. long, 1 mm. wide, 
compressed, 6-30-flowered, the rachilla straight, not winged, the achenes and sta- 
mens persistent after the glumes have fallen; scales 1-1.5 mm. long, oblong-ovate, 
obtuse, sometimes minutely mucronulate, reddish, thin, 3-nerved; achene rounded- 
trigonous, 0.5 mm. long, subglobose, stipitate, verrucose, scarcely apiculate. 

Kuekenthal refers all American material to subsp. juncoides, but 
this is separated from Old World plants by doubtfully distinct 
characters. 

Cyperus hermaphroditus (Jacq.) Standl. Contr. U. S. Nat. 
Herb. 18: 88. 1916. Carex hermaphrodita Jacq. Coll. Bot. 4: 174. 
1790. Mariscus Jacquinii HBK. Nov. Gen. & Sp. 1: 216. 1815. 
Cyperus alpinus Liebm. Vid. Selsk. Skrivt. V. 2: 215. 1851. Coyolito. 

Damp or wet thickets, fields, or forest, often in sand along streams 
or a weed in cultivated ground, ascending from sea level to about 



126 FIELD IANA: BOTANY, VOLUME 24 

2600 meters; Pete"n; Alta Verapaz; reported from Izabal; Zacapa; 
Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla; Sacatepe*quez; 
Chimaltenango; Suchitepe"quez ; Solola; Retalhuleu; Quezaltenango; 
San Marcos; Huehuetenango. Mexico and British Honduras to Pan- 
ama; South America. 

A glabrous perennial with short rhizomes, the culms 15-75 cm. tall, 1-2 mm. 
thick at the apex, trigonous, smooth, often tuberous-thickened at the base; leaves 
3-9 to a culm, 3-7 mm. wide, flat, scaberulous on the margins, the sheaths purplish 
brown; bracts 5-8, unequal, 2-7 mm. wide; rays 12 or fewer, 12 cm. long or less, 
sometimes wanting, often branched, the spikes 1-3 cm. long, 1-1.5 cm. broad, 
dense or lax, usually cylindric; spikelets numerous, 5-10 mm. long, 1 mm. wide, 
oblong-lanceolate, somewhat tetragonous, usually divaricate, 3-7-flowered, the 
lowest sometimes reflexed, the rachilla winged; scales 2.5-3.5 mm. long, ovate- 
elliptic, appressed, thin, obtuse or submucronate, many-nerved, yellowish to golden 
brown; achene 1.2-1.8 mm. long, trigonous, oblong-ellipsoid, not stipitate, minutely 
apiculate. 

Cyperus humilis Kunth, Enum. PI. 2: 23. 1837. C. humilis var. 
elatior Britton, Bull. Torrey Club 15: 99. 1888. 

Wet thickets, marshes, or sandy river beds, ascending from near 
sea level to about 1450 meters; Pete*n; Alta Verapaz; Zacapa. Mex- 
ico and British Honduras; West Indies; Peru. 

A glabrous annual with fibrous roots, the culms densely cespitose, 1-20 cm. 
tall, 1 mm. thick, smooth; leaves 1-2 to each culm, 20 cm. long or less, 1-2 mm. 
wide, roughened on the margins, the sheaths reddish brown; bracts 2-4, unequal, 
15 cm. long or less; inflorescence a single globose head 3-18 mm. broad, very dense; 
spikelets numerous, oblong-lanceolate, 3-6 mm. long, 1.5-2 mm. wide, compressed, 
8-18-flowered, the rachilla straight, not winged; scales about 1 mm. long, ovate, 
3-nerved, short-mucronate, the keel green, the sides whitish, hyaline; achene 0.8 
mm. long, trigonous-ovoid, acuminate, olive to chestnut-brown, transversely rug- 
ulose, substipitate. 

Cyperus imbricatus Retz. Obs. Bot. 5: 12. 1789. C. radiatus 
Vahl, Enum. PI. 2: 369. 1806. C. radiatus var. elongatus Boeckel. 
Linnaea 36: 319. 1870. 

Shallow water at edge of lake, forming large colonies, Lago de 
Atescatempa, Jutiapa, 500 meters; reported from mouth of Rio Polo- 
chic, Izabal. Mexico; Costa Rica; West Indies; South America; Old 
World tropics. 

Plants perennial, robust, glabrous, with very short, ligneous rhizomes, the 
culms about a meter tall, trigonous, smooth, leafy at the base; leaves shorter than 
culms, coriaceous, 4-8 mm. wide, the sheaths fuscous; bracts 3-5, equaling or longer 
than the inflorescence; rays of the umbel 6-8, compound, the spikes corymbose- 
fasciculate, sessile or subsessile, cylindric, dense, 1-3 cm. long, 4-8 mm. wide; 
spikelets numerous, crowded, suberect, oblong-lanceolate or linear-lanceolate, 3-6 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 127 

mm. long, about 1 mm. wide, subcompressed, 10-20-flowered, the rachilla narrowly 
winged; scales 1.2 mm. long, closely imbricate, broadly ovate, obtuse, membrana- 
ceous, pale ferruginous, 3-5-nerved, terminating in a short excurved mucro; achene 
half as long as the glume, ovoid or ellipsoid, compressed-trigonous, stramineous, 
lustrous, scarcely apiculate. 

Cyperus ischnos Schlecht. Bot. Zeit. 7: 99. 1849. Chan-sola 
(Huehuetenango) . 

Grassy slopes in pine forest, 250-2100 meters; Pete"n; Zacapa; 
Chiquimula; Santa Rosa; Huehuetenango. Mexico and British 
Honduras; northern South America. 

A slender perennial with short rhizomes, the culms separated, 20-75 cm. tall, 
1 mm. thick at the apex, smooth, often tuberous-thickened at the base; leaves 2-4 
to a culm, 10-40 cm. long, filiform, 1.5 mm. wide or less, rough-margined, the 
sheaths reddish brown; bracts 3-5 and 18 cm. long or less, filiform, the inflorescence 
consisting of 1-4 dense sessile heads 5-15 mm. broad, containing 5 to many spike- 
lets; spikelets oblong-lanceolate, 5-9 mm. long, 2 mm. wide, compressed, acute, 
4-12-flowered, the rachilla straight, narrowly winged; scales 2.2-2.8 mm. long, 
ovate or oblong-lanceolate, tridentate at the apex, 7-9-nerved, spreading when 
mature, the keel green, the sides white-hyaline to yellow; achene 1.2-1.5 mm. long, 
obovoid or oblong-ellipsoid, brown, puncticulate, not stipitate. 

Cyperus lanceolatus Poir. in Lam. Encycl. 7: 245. 1806. C. 
Humboldtianus Schult. in Roem. & Schult. Syst. 2: Mant. 100. 1824. 
C. Olfersianus Kunth, Enum. PI. 2: 10. 1837. C. lanceolatus var. 
compositus Presl, Reliq. Haenk. 1: 167. 1830. Tule. 

Wet fields or thickets, often along streams or lake shores or in 
marshes or ditches, ascending from sea level to about 2000 meters; 
Alta Verapaz; Baja Verapaz; Chiquimula; Jalapa; Escuintla; Gua- 
temala; Suchitepe"quez; Solola; Retalhuleu; Huehuetenango. South- 
ern United States to Mexico and British Honduras; West Indies and 
South America. 

A glabrous perennial with short rhizomes, the culms cespitose, 15-40 cm. tall, 
1 mm. thick at the apex, trigonous, smooth; leaves 2-4 to a culm, shorter than the 
culm, 1-2 mm. wide, flat, scaberulous on the margins, the sheaths reddish brown; 
bracts 2-4, unequal, 2-8 cm. long, longer than the inflorescence; rays of the umbel 
5 or fewer, often wanting, simple, bearing a cluster of 4-17 spikelets; spikelets lan- 
ceolate, 3-7 mm. long, 1.5-2 mm. wide, flat, 5-16-flowered, the rachilla straight or 
nearly so, not winged; scales 1.5 mm. long, broadly ovate, obtuse, 3-nerved, the 
keel green or brown, the sides hyaline; achene 1 mm. long, lenticular, obovate- 
ellipsoid, stipitate, the surface with hexagonal cells, reddish brown. 

Much of the Guatemalan material is referable to var. compositus 
Presl, in which the clusters of spikelets are pedunculate rather than 
sessile as in the typical form. 



128 FIELDIANA: BOTANY, VOLUME 24 

Cyperus lentiginosus Millsp. & Chase, Field Mus. Bot. 3: 74. 
1903. C. tennis var. lentiginosus Kuekenth. Pflanzenreich IV. 20: 
418. 1936. 

Damp thickets or fields, brushy slopes, or in sand along streams, 
at 1850 meters or less; Chiquimula; Jutiapa; Santa Rosa; Escuintla; 
Guatemala; Chimaltenango; Suchitepe"quez. Yucatan and British 
Honduras. 

A glabrous perennial with short rhizomes, the culms stout, 25-75 cm. tall, 
1.5 mm. thick at the apex, smooth; leaves 2-4 to a culm, flat, 2-6 mm. wide, rough- 
margined, the sheaths reddish purple; bracts 6-10 and 2-8 mm. wide, often much 
elongate; rays 6-16, simple or compound, the spikes 2-4 cm. long, 2-2.5 cm. broad, 
cylindric or oblong; spikelets distichous, linear-lanceolate, 10-13 mm. long, 1-1.5 
mm. wide, subterete, divaricate or the lower somewhat reflexed, 4-6-flowered, the 
rachilla straight, winged; scales 4-5 mm. long, oblong-lanceolate, short-mucronate, 
7-11-nerved, appressed or with somewhat spreading tips, clasping the achene, light 
brown spotted with dark purple; achene 2 mm. long, trigonous, obo void-oblong, 
brown, punctate, not stipitate. 






Cyperus ligularis L. PL Jam. Pugill. 3. 1759. Tul coyolillo. 

Wet fields or banks, especially in salt flats near the seacoast, as- 
cending to about 1170 meters (Lago de Amatitlan); Izabal; Chiqui- 
mula; Escuintla; Guatemala; Retalhuleu; San Marcos. Florida and 
Alabama and Mexico to British Honduras and Panama; West Indies; 
South America; Old World tropics. 

A coarse stout perennial with short thick rhizomes, the culms stout and stiff, 
30-90 cm. tall, trigonous, 2-5 mm. thick at the apex, papillose; leaves longer than 
the culm or shorter, 5-20 mm. wide, thick, septate-nodulose, papillose, very rough 
on the margins and costa, pale when dried, the sheaths reddish brown; bracts 5-12, 
unequal, 5-15 mm. wide; rays of the umbel 5-12, unequal, branched, usually short, 
the spikes 3-7 in a dense cluster, 1.5-2.5 cm. long, 10-12 mm. wide; spikelets very 
numerous, 2-6-flowered, turgid, subterete, 3-7 mm. long, 2-2.5 mm. wide, divari- 
cate or the lower reflexed, the rachilla zigzag, winged; scales 2-2.5 mm. long, ovate, 
apiculate, 9-11-nerved, brown or stramineous with reddish brown striations, lus- 
trous; achene 1.2-1.4 mm. long, trigonous, obo void-ellipsoid, brown, apiculate, 
puncticulate, substipitate. 

This isolated species grows most often in brackish soil, but not 
invariably so. In British Honduras it receives the name of "cutting- 
grass." 

Cyperus Lundellii O'Neill, Carnegie Inst. Wash. Publ. 522: 276. 
/. 2. 1940. 

Pete"n, the type from river bank, Subin River, Monte Santa 
Teresa, Lundell 2657; collected also at El Paso, Lundell 1475. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 129 

A glabrous perennial with short rhizomes, the culms 60-100 cm. tall, 2 mm. 
thick at the apex, triquetrous, scaberulous on the angles, obscurely septate-nodu- 
ilose; leaves 3-6 to a culm, 20-90 cm. long, 3-6 mm. wide, flat, rough-margined; 
ibracts 3-4 and 45 cm. long or less, 2-5 mm. wide; rays of the umbel 5-8, compound, 
the spikelets in clusters of 3-6 at the ends of the raylets; spikelets linear, acute, 
(6-13 mm. long, 2 mm. wide, turgid, 12-32-flowered, the rachilla straight, not 
winged; scales about 2 mm. long, suborbicular, acute and mucronulate, spreading, 
membranous, obsoletely 3-nerved, stramineous; achene 1 mm. long, trigonous, 
iovoid, black at maturity, acute, short-stipitate, conspicuously verrucose. 

Cyperus Luzulae (L.) Retz. Obs. Bot. 4: 11. 1786. Scirpus 
Luzulae L. Sp. PL ed. 2. 75. 1762. C. guatemalensisGandoger, Bull. 
Soc. Bot. France 66: 297. 1919 (type from Alta Verapaz, collected 
by Tuerckheim, probably at Cubilgliitz). Cebollin; Cebolla de rio; 
Sivac (Coban); Suchipaite (Huehuetenango) . 

Marshes and swamps, wet fields, or along streams, ranging from 
| sea level to about 1400 meters, most common at low elevations; 
'iPete'n; Alta Verapaz; Izabal; Retalhuleu; Suchitepe"quez; Huehuete- 
nango; Quezaltenango; San Marcos. Mexico and British Honduras 
to Panama; West Indies; South America. 

A glabrous perennial with short rhizomes, the culms usually cespitose, stout, 

1 20-50 cm. tall, obtusely trigonous; leaves 5-12 to a culm, 10-40 cm. long, 3-8 mm. 

wide, flat, rough-margined, the sheaths reddish brown; bracts 7-11, the rays of the 

umbel 6-12, the few spikes at the ends forming dense ovoid heads 6-15 mm. broad; 

spikelets ovate to oblong-ovate, 2.5-4 mm. long, 2 mm. wide, compressed, 6-16- 

flowered, the rachilla not winged; scales 1.2-1.5 mm. long, ovate or oblong-ovate, 

incurved, submucronate, obsoletely 3-nerved, white or yellowish; achene 1 mm. 

1 long, trigonous, linear-oblong, acuminate, scarcely stipitate, minutely puncticu- 

late, subapiculate. 

A weedy plant, common in many localities. The stems are much 
i| used as a substitute for twine. A decoction of the root is drunk in 
Huehuetenango with atol de maiz, often flavored with toasted seeds 
of sapote. 

Cyperus macrocephalus Liebm. Vid. Selsk. Skrivt. V. 2: 221. 
1851. C. oxycarioides Britton, Bull. Torrey Club 11: 86. 1884. 

Pete"n. Texas to Mexico and British Honduras. 

A perennial with short rhizomes, the stout culms more or less cespitose, 20-60 

I cm. tall, 1-2 mm. thick at the apex, trigonous, smooth; leaves 2-3 to a culm, 15-40 

; cm. long, 2-7 mm. wide, subcoriaceous, flat, rough-margined, the sheaths reddish 

brown; bracts 4-7, unequal, 4-10 cm. long; inflorescence densely congested and 

forming a solitary globose head 1-4 cm. in diameter; spikelets numerous, oblong to 

| linear-lanceolate, 5-12 mm. long, 1 mm. wide, 4-12-flowered, the rachilla disarticu- 

1 lating into 1-fruited joints, winged; scales 2 mm. long, ovate, sometimes mucronu- 



130 FIELDIANA: BOTANY, VOLUME 24 

late, obscurely 7-9-nerved, clasping the achene, stramineous to reddish brown, the 
margins white-hyaline; achene ellipsoid, obtusely trigonous, 1.2 mm. long, minutely 
puncticulate, short-stipitate, apiculate. 

The collection of J. D. Smith 1850 reported from mouth of Rio 
Polochic, Izabal, is to be referred to C. Eggersii. 

Cyperus manimae HBK. Nov. Gen. & Sp. 1: 209. 1815. C. 
asperrimus Liebm. Vid. Selsk. Skrivt. V. 2: 218. 1851. C. divergens 
HBK. op. cit. 208. C. manimae var. asperrimus Kuekenth. Pflan- 
zenreich IV. 20: 463. 1936. Chul, Chan-sola (Huehuetenango). 

Damp meadows or brushy slopes, sometimes in pine forest, often 
a weed in gardens or cornfields, 800-1900 meters; reported from 
Pet&i, probably in error; Escuintla; Chimaltenango; Sacatepe"quez; 
reported from Santa Rosa; Huehuetenango; Quezaltenango. South- 
western United States to Mexico and Costa Rica; mountains of 
South America. 

A perennial with short rhizomes, the culms 15-60 cm. tall, rather stout, smooth, 
leafy below; leaves shorter than the culms, 3-4 mm. wide, often scabrous, the 
sheaths brownish; bracts 5-8, longer than the inflorescence, this 3-6-radiate or con- 
tracted and almost head-like; the rays simple, the spikes ovate, 1.5-2 cm. long, 
dense; spikelets divergent, linear-lanceolate, acute, 5-6 mm. long, 1.5-2 mm. wide, 
6-10-flowered, the rachilla very narrowly winged; scales rather remote, ovate, 
rounded and short-mucronate at the apex, fulvous, 7-11-nerved, with white hya- 
line margins; achene three-fourths as long as the scale, trigonous, obo void-oblong, 
apiculate, brown or castaneous, lustrous. 

Var. asperrimus, distinguished by having somewhat scabrous 
culms, is represented by Guatemalan material. 

Cyperus Mutisii (HBK.) Griseb. Fl. Brit. W. Ind. 567. 1864. 
Mariscus Mutisii HBK. Nov. Gen. & Sp. 1: 216. 1815. C. martini- 
censis Boeckel. Cyp. Nov. 2: 38. 1890. C. Mutisii var. martinicensis 
Kuekenth. Pflanzenreich IV. 20: 485. 1936. C. Mutisii var. con- 
tractus Kuekenth. op. cit. 486. Pegapega (Chimaltenango) ; Coyolillo. 

Moist or wet fields or thickets, frequently in cultivated ground, 
often in sand along streams, ascending from about 200 meters to 1900 
meters, most frequent at middle or rather high elevations; Alta Vera- 
paz; Zacapa; Chiquimula; Jutiapa; Santa Rosa; Guatemala; Sacate- 
pe"quez; Chimaltenango; Quiche"; Huehuetenango. Mexico to British 
Honduras and Panama; West Indies and South America. 

A glabrous perennial with short ligneous rhizomes, the culms slender or rather 
stout, 30-100 cm. tall, 2-3 mm. thick at the apex, trigonous, usually smooth; leaves 
2-3 to a culm, 2-12 mm. wide, flat, rough-margined, the sheaths reddish brown; 
bracts 5-8, the rays of the umbel 7-10 (rarely none, in var. contractus) , with 1-6 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 131 

digitate spikes at the apex; terminal spikes 2-4.5 cm. long, 5-10 mm. wide, oblong- 
cylindric, the lateral ones shorter; spikelets ellipsoid, 3-5 mm. long, 1-2 mm. wide, 
divergent or reflexed, 1-5-flowered, the rachilla zigzag, winged; scales 2.2-3.2 mm. 
long, ovate or elliptic, obtuse, 9-13-nerved, sometimes mucronulate, yellow to red- 
dish or brown; achene 1.5-1.8 mm. long, broadly ellipsoid, brown, not stipitate, 
minutely apiculate, raised-puncticulate. 

This and the related species of subgenus Mariscus are rather 
poorly defined, and often it is difficult to decide to which supposed 
species a particular specimen should be referred. The group will 
require much additional study before the related plants can be clas- 
sified satisfactorily, if a satisfactory disposition of them ever is pos- 
sible. Kuekenthal refers Heyde & Lux 3542 from Dept. Santa Rosa 
to Cyperus subambiguus Kuekenth. (Pflanzenreich IV. 20: 527. 1936; 
Mariscus ambiguus Liebm. Vid. Selsk. Skrivt. V. 2: 231. 1851, not 
Cyperus ambiguus Liebm.). We have not seen the specimen thus 
reported, but C. subambiguus appears to be too close to C. Mutisii, 
and the definite characters by which it is to be separated are not 
obvious from Kuekenthal's account of the group. 

Cyperus niger Ruiz & Pavon, Fl. Peruv. 1: 47. 1798. C. melano- 
stachyus HBK. Nov. Gen. & Sp. 1: 207. 1815. Coyolillo; Juquillo. 

Marshes, wet fields or banks, along ditches, or often on sandbars 
along streams, 800-2500 meters; Alta Verapaz; Jalapa; Jutiapa; Gua- 
temala; Sacatepe*quez; Chimaltenango; Quiche"; Huehuetenango; 
Quezaltenango; San Marcos. Texas and New Mexico. Mexico to 
Panama, Peru, and Argentina. 

A small glabrous perennial with slender rhizomes, the culms usually cespitose, 
sometimes decumbent and rooting at the base, 10-30 cm. tall, smooth, with a few 
leaves at the base; leaves shorter than the culms, 1.5-2 mm. wide, the long sheaths 
brown; bracts 3, usually much longer than the inflorescence, this contracted and 
often head-like, 1-2 cm. broad, the rays 1-4 or none, bearing at the apex 3-6 spikes; 
spikelets crowded, spreading, oblong or linear-lanceolate, acute, 4-8 mm. long, 
2 mm. wide, compressed, 8-16-flowered, the rachilla straight, not winged; scales 

I densely imbricate, ovate-obtuse, dark red or dark castaneous, lustrous, 3-5-nerved; 

I achene two-thirds as long as the glume, elliptic-ovate, compressed-biconvex, brown 
and lustrous, finely striate, densely punctulate, apiculate. 

Kuekenthal and O'Neill recognize several varieties of the species. 
Cyperus niger var. capitatus (C. niger var. castaneus) differs from the 
species and the other varieties in the castaneous color of the scales 
and the conspicuously apiculate achenes which are almost as long as 
the scales. The species is usually easily distinguished from other 
species by its low stature and very dark spikelets. 



132 FIELD IANA: BOTANY, VOLUME 24 

Cyperus ochraceus Vahl, Enum. PL 2: 325. 1806. Tule. 

Bogs or wet meadows or thickets, ascending to 1900 meters; 
Pete"n; Zacapa; Chiquimula; Jalapa; Huehuetenango. Florida and 
Texas to Mexico, British Honduras, and Honduras; West Indies; 
South America. 

A glabrous perennial with short thick rhizomes, the culms 20-75 cm. tall, 
1-2 mm. thick at the apex, trigonous, smooth; leaves 6-12 to a culm, about equal- 
ing the culms, 2-6 mm. wide, minutely scaberulous on the margins, the sheaths 
purplish brown; bracts 5-8, very unequal, the rays of the umbel 6-12, short or 
elongate, often compound; spikelets oblong-lanceolate or linear, 5-20 mm. long, 
2-3 mm. wide, 16-30-flowered, forming globose heads, the rachilla straight, not 
winged; scales 1.5-2 mm. long, broadly ovate, not imbricate at maturity, cellular- 
reticulate, 3-nerved, falcate, stramineous to bright yellow or pale brown; achene 
ellipsoid to oval, 1.5 mm. long, conspicuously stipitate and acuminate-apiculate, 
purplish brown, white-reticulate. 

The Maya name reported from Yucatan is "mazcabzuuc." 

Cyperus odoratus L. Sp. PL 46. 1753. C. ferax L. Rich. Act. 
Soc. Hist. Nat. Paris 1: 106. 1792. C. Haenkei Presl, Reliq. Haenk. 
1: 172. 1830. Navajuela; Saichd (Coban, Kekchi). 

Usually in wet soil, thickets or waste ground, often along streams 
or ditches, marshes and wet fields, most common at low elevations, 
but ascending to about 1500 meters; Alta Verapaz; Izabal; Zacapa; 
Jutiapa; Santa Rosa; Escuintla; Guatemala; Retalhuleu; Quezalte- 
nango. United States to Mexico, British Honduras, and Panama; 
West Indies; South America; Old World tropics. 

A glabrous perennial with short rhizomes, the culms stout, 30-100 cm. tall, 
3-5 mm. thick at the apex, smooth; leaves 3-5 to a culm, 10-60 cm. long, 4-12 mm. 
wide, subcoriaceous and rather rigid, scaberulous on the margins, the sheaths stra- 
mineous or brown; bracts 6-8, elongate; rays of the umbel 6-12, short or elongate, 
compound, the spikes 2-3 cm. long and 1-2.5 cm. wide; spikelets mostly rather 
distant and lax, divergent or reflexed, linear, 5-15 mm. long, 1 mm. wide, sub- 
terete, 3-15-flowered, the rachilla readily breaking up into 1-fruited joints, 
winged; scales 2-3.5 mm. long, broadly ovate to ovate-elliptic, closely imbricate, 
appressed and enclosing the achenes, brownish or reddish-stramineous, obsoletely 
7-9-nerved; achene 1-1.5 mm. long, oblong or obovoid-oblong and trigonous, 
brown to black, substipitate, apiculate, minutely puncticulate. 

Called "coyolillo" in El Salvador and probably also in Guate- 
mala, the name alluding to a fanciful resemblance of the plant to a 
diminutive coyol palm (Acrocomia). This is a weedy plant and prob- 
ably the most abundant Cyperus species of Central America. 

Cyperus oxylepis Nees, Linnaea 9: 285. 1835, nomen; Steud. 
Syn. Cyper. 25. 1855. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 133 

In marshes, on sand, or in salt meadows of the Pacific seashore; 
Retalhuleu (Champerico) ; San Marcos (Ocos). West Indies and 
South America. 

A glabrous perennial with short ligneous rhizomes, the culms cespitose, 15- 
30 cm. tall or more, obscurely trigonous, glutinous as also the leaves; leaves few, 
about equaling the culms, narrow, coriaceous, obsoletely septate-nodulose, rough- 
margined, involute, the sheaths stramineous, or purplish at the base; bracts 3, 
foliaceous, much exceeding the inflorescence; rays 3-6, simple or compound, 
unequal, 15 cm. long or less, the spikelets several, forming a hemispheric head 
1.5-2 cm. in diameter; spikelets linear-lanceolate, acute, 10-20 mm. long, 3 mm. 
wide, subcompressed, 12-20-flowered, the slender rachilla not winged; scales rather 
remote, chartaceous, oblong-elliptic, patulous at the apex, the keel bright green, 
excurrent as an excurved mucro, the sides yellowish or brown, lustrous; achene 
three-fourths as long as the glume, oblong, attenuate at each end, trigonous, 
black at maturity, apiculate, white-punctulate. 

Cyperus pallens (Liebm.) Standl. & Steyerm., comb. nov. 
Mariscus pallens Liebm. Vid. Selsk. Skrivt. V. 2: 230. 1851. M. 
Haenkei Presl, Reliq. Haenk. 1: 181. 1830, not Cyperus Haenkei 
Presl. C. regiomontanus Britton, Contr. U. S. Nat. Herb. 1: 362. 
1895. C. Pittieri Boeckel. Allgem. Bot. Zeit. 2: 19. 1896. C. regiomon- 
tanus var. pallens (Liebm.) Kuekenth. Pflanzenreich IV. 20: 527. 
1936. C. regiomontanus var. Pittieri (Boeck.) Kuekenth. loc. cit. 

Reported by Kuekenthal from Huehuetenango, Jacaltenango, 
Seler 2860. California and western Mexico; British Honduras; Costa 
Rica; Venezuela; Peru. 

A perennial with short rhizomes, the culms cespitose, slender, 10-30 cm. tall, 
acutely angulate, smooth, thickened at the base; leaves equaling the culms, 1.5- 
2 mm. wide, flat, the sheaths dark purplish; bracts 3-5, much exceeding the 
nflorescence, the rays of the umbel 3-5, simple, very short, the spikes ovoid or 
oblong-cylindric, 8-14 mm. long, 6-8 mm. wide, dense; spikelets divaricate, linear- 
oblong, 2.5 mm. long, 1-2-flowered, the rachilla winged; scales oblong-elliptic, 
subobtuse, the keel green, the sides pale ferruginous, 9-nerved; achene three- 
xmrths as long as the scale, trigonous, slightly curved, apiculate. 

The single British Honduras collection cited by Kuekenthal is 
referred by O'Neill to C. flavus. The proper disposition of C. pallens 
of which we have seen no Guatemalan material is decidedly un- 
certain, and it may not be a distinct species. Kuekenthal uses for it 
the name C. regiomontanus, under which C. pallens is treated as a 
variety, a course in violation of current rules of nomenclature. 

Cyperus panamensis (Clarke) Britton ex Standl. Journ. Wash. 
Acad. Sci. 15: 457. 1925. Mariscus panamensis Clarke, Kew Bull. 
Add. Ser. 8: 15. 1908. 



134 FIELDIANA: BOTANY, VOLUME 24 

Wet open ground, 200-300 meters, Escuintla; Zacapa. Panama; 
Ecuador and Peru. 

Plants annual, the culms cespitose, 20-30 cm. tall, triquetrous, rather stout, 
smooth, leafy below; leaves shorter than the culms, 3-5 mm. wide, flat, the sheaths 
dark brown at the base; bracts 4-5, longer than the inflorescence; rays of the umbel 
3-8, simple, 5 cm. long or less, the spikes ovate-cylindric or broadly ovate, 2 cm. 
broad; spikelets numerous and usually dense, linear-oblong, acute, 7-10 mm. long, 
1.5 mm. wide, obsoletely quadrangular, 3-8-flowered, spreading or divergent, the 
rachilla broadly winged; scales rather remote, oblong-elliptic, obtuse, the costa 
ex current as a long mucro, the sides fulvous, 7-9-nerved; achene two-thirds as long 
as the glume, broadly oblong, slightly curved, trigonous, blackish, densely punctu- 
late, short-apiculate. 

Called "coyolillo" in El Salvador. 

Cyperus piceus Liebm. Vid. Selsk. Skrivt. V. 2: 200. 1851. 

Reported by Kuekenthal from vicinity of Coban, Alta Verapaz, 
1350 meters, Tuerckheim 1933. Southern Mexico to Panama; South 
America. 

A glabrous annual, the culms cespitose, slender, 10-30 cm. tall, smooth, 
sparsely leafy below; leaves shorter than the culms, very narrow, flat, the sheaths 
purplish; bracts 2-3, usually longer than the inflorescence; rays of the umbel 3-4, 
simple, the rays 4 cm. long or less, with 3-6 clustered spikelets at the tips; spike- 
lets linear-oblong, 6-12 mm. long, 2 mm. wide, compressed, 10-20-flowered, the 
rachilla straight; scales rather densely imbricate, ovate, obtuse, castaneous, the 
margins white-hyaline, 3-costate; achene half as long as the glume, broadly obo- 
vate, biconvex, scarcely 1 mm. long, blackish, lustrous, transversely zonate, short- 
stipitate, apiculate. 

Cyperus polystachyos Rottb. Desc. & Icon. 39. 1773. C. odo- 
ratus Auct., not L. C. paniculatus Rottb. op. cit. 40. C. fugax Liebm. 
Vid. Selsk. Skrivt. V. 2: 196. 1851. 

Wet or moist fields or plains, 200 meters or less; Izabal; Zacapa; 
reported from Santa Rosa at 900 meters. Mexico to British Hon- 
duras and Panama; West Indies; South America; Old World tropics. ' 

Perennial, with short rhizomes, the culms 20-50 cm. tall, slender, 1 mm. thick 
at the apex, smooth; leaves 2-5 to a culm, shorter than the culms, 2-4 mm. wide, 
coriaceous, rough-margined, the sheaths reddish brown; bracts 3-6, unequal, longer 
than the inflorescence; rays of the umbel 4-6, compound, unequal, 4 cm. long or 
less; spikelets 7-10 mm. long, 1.5 mm. wide, compressed, ascending, 13-40-flow- 
ered, the rachilla zigzag, winged; scales 1.5 mm. long, oval, obtuse, 3-nerved, 
stramineous, with hyaline margins; achene 1 mm. long, lenticular, obovate-oblong, 
the surface with hexagonal cells, short-stipitate, short-apiculate, brown to black, 
densely puncticulate. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 135 

Called "pelillo" in El Salvador. Kuekenthal recognizes several 
varieties and forms of the species, but they seem scarcely worth 
nomenclatural recognition. 

Cyperus prolixus HBK. Nov. Gen. & Sp. 1: 206. 1816. 

Marshes and open swamps, ranging from sea level to about 1500 
meters; Alta Verapaz; Izabal; Guatemala; reported by Hemsley 
from San Geronimo and Volcan de Fuego; collected by Godman and 
Salvin. Southern Mexico; Costa Rica and Panama; South America. 

Plants perennial, large and coarse, with thick rhizomes, the culms commonly 
1-1.5 meters tall, obtusely trigonous, spongy-thickened at the base, leafy below; 
leaves equaling or longer than the culms, 8-20 mm. wide, flat, septate-nodulose, 
rough-margined; bracts 6-10, longer than the inflorescence; umbels decompound 
or even thrice branched, lax, large, with numerous rays, these as much as 30 cm. 
long; spikes rather dense, with numerous spikelets, oblong-elliptic; spikelets sub- 
erect, linear or linear-lanceolate, 15-20 mm. long, 1.5-2 mm. wide, acute, com- 
pressed, 10-14-flowered, the very slender rachilla flexuous, winged; scales rather 
remote, patulous at the apex in fruit, oblong-elliptic, obtuse, 4 mm. long, often 
short-mucronate, dirty-stramineous, 5-7-flowered, ferruginous-lineolate between 
the nerves; achene one-half to three-fifths as long as the glume, trigonous, narrowly 
oblong, dark brown, lustrous, densely punctulate, short-apiculate. 

The plant is abundant in many places in the lowlands of Izabal, 
where it forms almost pure stands of considerable extent. In general 
appearance it is much like the papyrus of the Nile. 

Cyperus pseudovegetus Steud. Syn. Cyp. 24. 1855. C. pseu- 
dovegetus var. megalanthus Kuekenth. Pflanzenreich IV. 20: 176. 1936 
(type from Cubilgiiitz, Alta Verapaz, 350 meters, Tuerckheim 975}. 

Stream banks, Pete"n and Alta Verapaz; no Guatemalan speci- 
mens have been seen by the writers. United States and Mexico. 

A glabrous perennial with short ligneous rhizomes, the culms 40-75 cm. tall, 
trigonous, smooth, leafy below; leaves equaling the culms, 2-10 mm. wide, flat, 
scarcely septate-nodulose; bracts 4-6, very long, the rays of the umbel about 8, 
simple or compound, 6 cm. long or less, usually bearing 3-6 subglobose heads of 
spikelets more or less contracted into a single head; spikelets numerous, ovate, 
densely congested into a dense lobate head, 4-7 mm. long, 3-4 mm. wide, 10-20- 
flowered, the rachilla not winged; scales densely imbricate, finally patulous at the 
apex, oblong-lanceolate, ferruginous to greenish, the costa excurrent as a conspic- 
uous excurved mucro; achene 1.2-1.8 mm. long, obscurely trigonous, linear, pur- 
alish brown, conspicuously stipitate, apiculate, minutely puncticulate. 

This species is very closely related to C. surinamensis. The Gua- 
:emalan collections are referable to var. megalanthus, distinguished 
}y having wider leaves and somewhat larger heads of spikelets than 
the typical form. 



136 FIELDIANA: BOTANY, VOLUME 24 

Cyperus rivularis Kunth, Enum. PL 2: 6. 1837. C. lagunelto 
Steud. Syn. Cyp. 5. 1855. C. rivularis subsp. lagunetto Kuekenth. 
Pflanzenreich IV. 20: 373. 1936. C. rivularis var. lagunetto (Steud.) 
O'Neill, Rh. 44: 86. 1942. 

Sandy stream banks, 850-1400 meters; Jutiapa; Alta Verapaz; 
Huehuetenango. United States and Mexico; Honduras; Costa Rica; 
South America. 

A glabrous annual with fibrous roots, the culms cespitose, slender, 5-60 cm. 
tall, smooth, leafy at the base; leaves mostly longer than the culms, setaceous, soft 
and weak, the sheaths brown; bracts 3-4, most of them longer than the inflores- 
cence; rays of the umbel 2-5, simple, 2 cm. long or less, bearing at the apex a cluster 
of 4-8 spikelets; spikelets rather densely subspicate, divergent, oblong, 6-12 mm. 
long, 2-2.5 mm. wide, obtuse, compressed, 10-24-flowered, the rachilla straight, 
not winged; scales densely imbricate, oblong-ovate, obtuse, fuscous or variegated 
with chestnut, lustrous, green-carinate, 3-nerved; achene half as long as the glume, 
obovate, compressed-biconvex, almost black, lustrous, short-apiculate. 

The Guatemalan and all tropical American material is referred by 
Kuekenthal to subsp. lagunetto, which he separates upon the basis of 
often taller culms, flat and broader leaves, and laxly spicate leaflets, : 
but O'Neill has reduced the subspecific rank to varietal category, 
separating the variety lagunetto on the basis of differently shaped 
achenes and sometimes 3 stamens. 

Cyperus rotundus L. Sp. PI. 45. 1753. Coyolillo. Figure 25. 

Wet fields or along streams, sometimes in lawns or pastures, 
chiefly at low elevations but ascending to possibly 1500 meters; 
Izabal; Zacapa; Guatemala. United States and Mexico to British 
Honduras and Panama; West Indies; South America; Old World 
tropics. 

A low perennial with long stolons ending in small tubers, the culms slender, 
15-50 cm. tall, 1-1.5 mm. thick at the apex, triquetrous, smooth; leaves 5-12 to a 
culm, about equaling the culms, flat, 3-6 mm. wide, smooth on the margins, the 
sheaths reddish brown; bracts 2-4, longer or shorter than the inflorescence; rays 
of the umbel 3-8, usually simple, sometimes compound, 9 cm. long or less; spikelets 
2-12 in each spike, lax, linear, compressed, 4-40 mm. long, 1-2.5 mm. wide, 12-36- 
flowered, the rachilla winged; scales imbricate, ovate, obtuse, thin, 3-3.5 mm. long, 
reddish brown, the keel green, 7-nerved, scarcely mucronate; achene 1.5-2 mm. 
long, obovoid or ellipsoid, black and shining, scarcely apiculate, minutely punc- 
ticulate. 

On the North Coast this species is often a weed in flower beds, 
gardens, and other cultivated ground 




FIG. 25. Cyperus rotundus. a, Spikelet (X 5). 6, Habit (X l /$). 



137 



138 FIELDIANA: BOTANY, VOLUME 24 

Cyperus semiochraceus Boeckel. Flora 61: 29. 1878. C.Bour- 
gaei Clarke ex Lundell, Carnegie Inst. Wash. Publ. 436: 287. 1934, 
nomen nudum. 

Pete"n (Uaxactun). Southern Mexico to British Honduras. 

A stout coarse perennial with short thick ligneous rhizomes, the culms 1-1.5 
meters tall, 8 mm. thick at the apex, obtusely trigonous, smooth; leaves coriaceous, 
shorter than the culms, 1.5-2 cm. wide, serrulate on the margins, the sheaths red- 
dish purple; bracts 5-12, much exceeding the inflorescence; rays 7-12, compound, 
50 cm. long or less, the spikes partly sessile and partly pedunculate, fasciculate, 
long-cylindric, 2-6 cm. long, 1-2 cm. thick, dense; spikelets divaricate or ascending, 
linear-oblong or linear, 3-15 mm. long, 1 mm. wide, subcompressed, 8-10-flowered, 
the rachilla winged; scales closely imbricate, 1.3-1.8 mm. long, lustrous, ovate or 
obovate, sometimes mucronulate, obscurely nerved, brown, with yellow-hyaline 
margins; achene 0.8 mm. long, compressed-trigonous, oval or ovoid, short-stipitate, 
scarcely apiculate, pale brown. 

Called "zacate cortador" in Campeche. 

Cyperus seslerioides HBK. Nov. Gen. & Sp. 1: 209. 1815. 

Reported by Kuekenthal from Quezaltenango, Bernoulli & Carlo 
1003; Jalapa; reported from Santa Rosa; Huehuetenango. Mexico; 
Venezuela. 

A low perennial with short rhizomes, the culms cespitose, slender, 5-16 cm. tall, 
obscurely trigonous, smooth, bulbous-thickened at the base; leaves few, about 
equaling the culms or shorter, very narrow, complicate, rather stiff, the sheaths 
brown; bracts 3, longer than the inflorescence, reflexed, the inflorescence contracted 
into a single subglobose head 1 cm. or less in diameter; spikelets numerous, oblong- 
lanceolate, subobtuse, compressed, 3-4 mm. long, 2-2.5 mm. wide, 10-12-flow- 
ered, the rachilla straight, not winged; scales closely imbricate, membranaceous, 
finally patulous at the apex, lance-ovate, 1.5 mm. long, pale stramineous or 
fulvous, 3-nerved, acuminate and terminating in an excurved mucro; achene one- 
third as long as the glume, rounded-ovate, trigonous, obtuse, dark brown, densely 
punctulate. 

Cyperus simplex HBK. Nov. Gen. & Sp. 1: 207. 1816. 

Wet thickets or forest, 1150 meters or less; Izabal (Quirigua); 
Huehuetenango. Southern Mexico; Costa Rica; widely dispersed in 
South America. 

A slender perennial with short rhizomes, the culms few, cespitose, 4-15 cm. 
tall, smooth, leafy below; leaves much longer than the culms, 3-6 mm. wide, flat, 
thin, the sheaths purplish; bracts 4-10, equaling or longer than the rays; umbel 
simple, the rays 5-10, very unequal, 5-20 cm. long, bearing apical clusters of 
usually 1-3 digitate spikelets; spikelets linear-oblong, acutish, 6-16 mm. long, 
2 mm. wide, subcompressed, 12-36-flowered, the rachilla flexuous, not winged; 
scales lax, patulous at the apex, membranaceous, ovate, incurved, articulate at 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 139 

the base, scabrous-carinate, obscurely 5-7-nerved, pale brownish, sometimes red- 
lineolate; achene two-fifths as long as the glume, obovoid, trigonous, truncate at 
the apex and apiculate, pale rufous, densely white-puncticulate, lustrous. 

In general appearance similar to C. diffusus, but easily recognized 
by the very short culms and the much longer rays of the umbel. 
Apparently rare in Guatemala, as in other parts of Central America. 

Cyperus surinamensis Rottb. Descr. & Icon. 35. 1773. C. suri- 
namensis var. lutescens Boeckel. Linnaea 35: 555. 1868. Coyolillo. 

Wet fields and thickets, swamps, stream and lake margins, often 
in sand flats along streams, chiefly at less than 1000 meters, rarely 
ascending to 1800 meters; Izabal; Zacapa; Chiquimula; Jalapa; Juti- 
apa; Santa Rosa; Escuintla; Guatemala; reported from Sacatepe"- 
quez; Chimaltenango; Huehuetenango; San Marcos. Florida to 
Texas, Mexico, British Honduras, and Panama; West Indies; South 
America. 

A glabrous perennial with short thick rhizomes, the culms stout, 10-60 cm. 
tall, 1 mm. thick at the apex, trigonous, downwardly scabrous on the angles; leaves 
2-6 to a culm, about equaling the culms, 1-3 mm. wide, flat or conduplicate, bright 
green, rough-margined, the sheaths purplish or brownish; bracts 5-7, very unequal, 
the rays of the umbel 5-7 and 7 cm. long or less, sometimes compound; spikelets 
congested in dense heads, numerous, oblong or linear, 3-14 mm. long, 2 mm. wide, 
strongly compressed, the rachilla not winged; scales 1-1.5 mm. long, ovate, acute 
or minutely apiculate, cellular-reticulate, stramineous, 3-nerved; achene 0.6 mm. 
long, narrowly ovoid-oblong, obtusely trigonous, faintly transverse-rugulose, 
minutely puncticulate, short-apiculate, reddish brown. 

Cyperus tenerrimus Presl, Reliq. Haenk. 1: 166. 1828. 

Grassy banks or fields, wet sand along streams, 200-1600 meters; 
Zacapa; Jutiapa; Suchitepe"quez; Santa Rosa; Escuintla; Guatemala; 
Quiche" ; Huehuetenango. Mexico to Costa Rica; Colombia. 

A low perennial with very short rhizomes, the culms very slender, cespitose, 
5-15 cm. tall, smooth, thickened at the base; leaves shorter or longer than the 
culms, 1 mm. wide, the sheaths brown; bracts 6-8, very long and spreading, finally 
reflexed; inflorescence consisting of a single dense head 5-15 mm. in diameter; 
spikelets numerous, elliptic or ovate, 3-5 mm. long, 2 mm. wide, compressed, 
6-16-flowered, acutish, the rachilla straight, not winged; glumes densely imbricate, 
finally patulous at the apex, incurved, oblong-elliptic, navicular and carinate, 
truncate-retuse at the apex, dirty white, 5-nerved; achene two-thirds as long as 
the glume, narrowly oblong, trigonous, apiculate, blackish, punctulate. 

Called "pelillo" in El Salvador and "tule" in Oaxaca. 

Cyperus tenuis Swartz, Prodr. Veg. Ind. Occ. 20. 1788. C. cara- 
casanus Kunth, Enum. PI. 2: 86. 1837. Zumbidor (Santa Rosa). 



140 FIELDIANA: BOTANY, VOLUME 24 

Wet or damp fields or thickets, often on sand or gravel bars, 1400 
meters or less, chiefly at low elevations; Pete"n; Alta Verapaz; Izabal; 
Zacapa; Santa Rosa; Escuintla; Suchitepe'quez; Quezaltenango; San 
Marcos. Mexico and British Honduras to Panama; West Indies; 
South America; west Africa. 

A glabrous perennial with short rhizomes, the culms cespitose, 10-50 cm. tall, 
1-1.5 mm. thick at the apex, trigonous, smooth; leaves 3-5 to a culm, 5-30 cm. 
long, 2-3 mm. wide, flat, rough-margined, the sheaths reddish purple; bracts 5-9, 
longer than the inflorescence; rays of the umbel 5-12, simple, 9 cm. long or less, 
the spikes 1.5-2 cm. long, 1-2.5 cm. wide; spikelets rather dense or remote, linear, 
5-15 mm. long, 3-8-flowered, the rachilla winged; scales 3-3.5 mm. long, oblong- 
elliptic, obtuse, mucronulate, closely appressed, 7-11-nerved, greenish white to 
brown, with hyaline margins; achene 1.5-2 mm. long, trigonous, oblong-ellipsoid, 
brown, somewhat falcate, substipitate, scarcely apiculate, punctate, dull. 

Called "coyolillo" in El Salvador. 

Cyperus uncinulatus Schrad. ex Nees in Mart. Fl. Bras. 2, 
pt. 1:23. 1842. 

Moist or dry, open or brushy plains and hillsides, about 200 
meters; Zacapa (collected several times in the vicinity of Zacapa). 
Southern and western South America. 

Plants with short rhizomes, the culms very slender, 5-12 cm. high, compressed- 
trigonous, smooth, bulbous-thickened at the base; leaves often equaling or longer 
than the culms, scarcely 1 mm. wide; bracts 3-4, much longer than the inflores- 
cence, spreading or reflexed; inflorescence capitate, subglobose, 5-10 mm. in diam- 
eter, very dense, with 7 or much more numerous spikelets; spikelets oblong, 4-6 
mm. long, 2-2.5 mm. wide, compressed, 10-20-flowered, the rachilla not winged; 
scales dense, strongly outcurved above, oblong-ovate, ferruginous or sordid-stra- 
mineous, cuspidate with a long excurved mucro; stamen 1; achene obovate-oblong 
or obovate, trigonous, rufous, densely punctulate, short-apiculate; style short, 3-fid. 

Cyperus unioloides R. Br. Prodr. Nov. Roll. 216. 1810. C. 
bromoides Willd. ex Link, Jahrb. 3: 85. 1820. 

In marshes, 300-1700 meters; Alta Verapaz; Jalapa; Santa Rosa; 
reported from Sacatepe"quez; Huehuetenango. California to Mexico 
and British Honduras; West Indies; South America. Tropics of 
Old World. 

A perennial with very short rhizomes, the culms cespitose, 20-75 cm. tall, 
1.5 mm. thick at the apex, rather stout, smooth; leaves 2-4 to a culm, shorter than 
the culms, 1-4 mm. wide, rough-margined, the sheaths reddish brown; bracts 2-4, 
unequal, longer than the inflorescence; rays 2-5, simple, unequal, 15 cm. long or 
less; spikelets 4-16 in each spike, lax, 10-15 mm. long, compressed, 3-4 mm. wide, 
12-24-flowered, the rachilla zigzag; scales 3.8 mm. long, ovate, acute, 3-costate, 
yellow and red-lineolate, the margins white-hyaline; achene 1 mm. long, lenticular, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 141 

obovate, grayish to black, puncticulate, sessile, short-apiculate, conspicuously 
cellular. 

Cyperus virens Michx. Fl. Bor. Amer. 1: 28. 1803. C. virens 
var. glaucopattidus Boeckel. Linnaea 35: 554. 1868. Sacabasto. 

Wet soil in quebradas or on lake shores, 1400-2400 meters; Alta 
Verapaz; Jalapa; Huehuetenango. United States and Mexico; Costa 
Rica; West Indies; South America. 

A coarse perennial with short thick rhizomes, the stout culms 30-90 cm. tall, 
triquetrous, rough-angled, leafy near the base; leaves equaling or shorter than the 
culms, 6-10 mm. wide, septate-nodulose, rough-margined, the sheaths purplish; 
bracts 5-7, very long, the rays of the umbel compound, unequal, 6 cm. long or less; 
spikelets forming globose heads of 6 or more, ovate-lanceolate or oblong-lanceolate, 
acute, 6-8 mm. long, 2.5-3 mm. wide, compressed, 10-20-flowered, the rachilla 
straight, not winged; scales dense, finally patulous, oblong-ovate, subacute, cellu- 
lar-reticulate, pale rufous or yellowish-stramineous, 3-nerved, sometimes minutely 
mucronate; achene about three-fifths as long as the glume, oblong or oblong-elliptic, 
trigonous, rufescent, dull, minutely punctulate, conspicuously stipitate, apiculate. 

DICHROMENA Michaux 

Plants usually perennial and with rhizomes, the leaves narrow; spikelets 
crowded in a terminal head on the leafless culm, involucrate by the leaflike bracts, 
these often tinged with white within near the base; spikelets compressed, with few 
or many flowers, the scales spirally imbricate, several of them empty or with im- 
perfect flowers; perianth none; stamens 3; style 2-cleft, the branches subulate; 
achene lenticular, transversely rugose, crowned with the broad persistent style 
base (tubercle). 

About 20 species, in North and South America, chiefly in the 
tropics. Only the following species are known from Central America. 

Leaves 5-12 mm. wide; bracts green throughout D. Watsoni 

Leaves 2-4 mm. wide; bracts often white at the base. 

Bracts glabrous, not ciliate; plants with elongate rhizomes often 10-30 cm. long. 

D. color ata 

Bracts ciliate at the base and sometimes also pilose; rhizomes none or short, 
rarely as much as 2 cm. long. 

Plants with fibrous roots; bracts green throughout; spikelets 1-3, cinnamon- 
brown when dry D. radicans 

Plants with short rhizomes; bracts white within near the base; spikelets 3-15, 
white when dry D. ciliata 

Dichromena ciliata Vahl, Enum. PI. 2: 240. 1806. D. nervosa 
Vahl, op. cit. 241. 1806. Junquillo (Pete"n, fide Lundell). 

Grassy open places, sometimes in pine forest; Pete"n; Alta Vera- 
paz; Izabal; Zacapa; Jalapa; Jutiapa; Huehuetenango; at 2000 



142 FIELDIANA: BOTANY, VOLUME 24 

meters or less; reported also from other departments, but the col- 
lections were not seen and their determinations therefore are ques- 
tionable. Southern Mexico and British Honduras to Panama, 
southward through much of South America; West Indies. 

Plants often in small dense clumps, usually 20-40 cm. high, the rhizomes very 
short, the culms slender and wiry, sometimes hirsute at the apex; leaves 10-30 cm. 
long, mostly 1-3 mm. wide, glabrous or often pilose on the margins or beneath, 
1-nerved, the sheaths brownish; bracts 4-6, white at the base; spikelets 3-15 and 
5-10 mm. long, usually bright white, each containing 3-10 achenes; scales 3-5 mm. 
long, ovate or lanceolate; achene 1.2 mm. long, suborbicular or ovoid, dark brown, 
transversely rugose, biconvex, bearing at the apex a wide tubercle. 

Although abundant in some parts of Central America, in Guate- 
mala this species appears to be rather rare and of only sporadic 
occurrence. 

Dichromena colorata (L.) Hitchcock, Ann. Kept. Mo. Bot. 
Card. 4: 141. 1893. Schoenus coloratus L. Sp. PI. 43. 1753. D. leu- 
cocephala Michx. Fl. Bor. Amer. 1: 37. 1803. 

Wet soil, often in savannas, 1600 meters or less; Pete"n; Zacapa. 
Eastern and southern United States through Mexico to British 
Honduras. 

Plants 30-50 cm. tall, the rhizomes often 10-30 cm. long, very slender, the 
slender culms erect; leaves 10-30 cm. long, 1-3 mm. wide, flat or conduplicate, 
the sheaths pale green or brownish; bracts 4-6 and 1-12 cm. long, similar to the 
leaves, often reflexed, white at and above the base, glabrous; spikelets numerous, 
sometimes 30 or more, 6-8 mm. long, each containing 6-10 achenes; scales broadly 
ovate-lanceolate, 3-4 mm. long, subacute; achene about 0.8 mm. long, truncate at 
the apex, with a flat tubercle, orbicular to obovate, transversely rugulose. 

Dichromena radicans Schlecht. & Cham. Linnaea 6: 38. 1831. 
Coyolillo; Punac (Coban, Kekchi). 

Usually in wet soil, frequently in ditches or marshes, often in wet 
or moist forest or thickets, sometimes in open places, frequently a 
weedy plant, ascending to about 1500 meters but most plentiful at 
low elevations; Alta Verapaz; Baja Verapaz; Izabal; Chiquimula; 
Santa Rosa; Guatemala; Suchitepe"quez; Retalhuleu; Huehuete- 
nango; Quezaltenango; San Marcos. Mexico to British Honduras 
and Panama; West Indies and South America. 

Plants perennial and densely cespitose, forming small or large clumps, the 
rhizomes very short or none, the culms 20-40 cm. tall, glabrous; leaves 10-30 cm. 
long, usually 1-3 mm. wide, flat, 5-9-nerved, ciliolate, the sheaths green or whitish, 
pubescent; bracts 2-5, 7 cm. long or less, commonly ciliate and also pilose near the 
base, green throughout; spikelets 1-5, lanceolate, acuminate, 8-10 mm. long, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 143 

brownish, at least when dry; scales 4-6 mm. long, ovate or lanceolate; achene 
lenticular, orbicular, 0.8 mm. long, smooth and lustrous, broadly stipitate, cream- 
colored, bearing a short tubercle. 

Dichromena Watsoni Britton, Bull. Torrey Club 15: 101. 1888. 
Figure 26. 

Type collected in Guatemala by Sereno Watson, probably in Iza- 
bal; wet forest, usually at low elevations, in western Guatemala 
ascending to about 1500 meters; Izabal; Alta Verapaz; Huehuete- 
nango; San Marcos (Volcan de Tajumulco). British Honduras to 
Panama. 

Plants rather stout, as much as 60 cm. high, with short or no rhizomes; cauline 
leaves 15-25 cm. long, 5-12 mm. wide, about 7-nerved, glabrous, the sheaths green; 
bracts about 9, or fewer, similar to the cauline leaves; spikelets about 9, acute, 
12-15 mm. long, greenish white; scales ovate-lanceolate, acute and mucronate, 
glabrous; achene obovate, transversely rugose, with a broad depressed tubercle. 

A relatively rare species of which but few collections are known, 
these all Central American. 



ELEOCHARIS R. Brown 

Reference: H. K. Svenson, Monographic studies in the genus 
Eleocharis, Rhodora 31: 121-135; 152-163; 167-191; 199-219; 224- 
242. 1929; 34: 193-203; 215-227. 1932; 35: 377-389. 1934; 39: 210- 
231; 236-273. 1937; 41: 1-19; 43-77; 90-110. 1939; Scirpeae, 
N. Amer. FI. 18: 509-540. 1957. 

Glabrous annuals or perennials; culms simple, terete to trigonous, quadrangu- 
lar, or compressed; leaves reduced to bladeless basal sheaths; spikelets terminal, 
solitary, erect, containing several to many flowers; scales concave, spirally imbri- 
cate or subdistichous; perianth of 6 or 8 bristles, these mostly retrorse-barbellate; 
stamens 2-3; style 2-3-cleft; achene lenticular or trigonous; style base enlarged 
and persistent as a tubercle upon the apex of the achene. 

About 150 species, widely distributed in almost all parts of the 
earth. A few additional species may occur in southern Central 
America. The following treatment is adapted from that by Svenson. 
The name is sometimes written Heleocharis. Although Dr. Svenson 
studied and identified our Guatemalan material in 1939 and 1940, 
the data were not incorporated, for some reason, in the distributional 
ranges published in the latest treatment in North American Flora. 
In the present treatment, it will be noted, there are cited for Guate- 
mala and British Honduras, as well as for other Central American 
countries, numerous species whose distribution was either omitted or 
incompletely cited in the North American Flora for those countries. 




FIG. 26. Dichromena Watsoni. a, Habit (X 1 A). b, Spikelet (X 4). 



144 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 145 

Scales firm, somewhat indurate, yellowish, scarcely carinate or nerved. Achenes 
lenticular; style 2-3-fid; culms usually stout and thick. 

Culms septate E. interstincta 

Culms not septate. 

Culms terete E. cellulosa 

Culms ungulate. 

Culms slender, 1-2 mm. thick E. plicarhachis 

Culms stout, 2-5 mm. thick. 

Achene constricted below the summit into a neck half as wide as the 

achene E. fistulosa 

Achene not constricted, gradually prolonged into a cellular beak. 

E. mutata 

Scales thin, conspicuously keeled or nerved. Achenes lenticular or trigonous; 
culms stout, or often very slender. 

Culms septate, sometimes only faintly so, 1-10 mm. thick. Plants perennial. 

Culms 1-2 mm. thick; spikelets 8-15 mm. long, much thicker than the culm. 

E. montana 

Culms 3-10 mm. thick; spikelets 15-40 mm. long, not thicker than the culm. 

E. elegans 

Culms not septate, very slender, less than 5 mm. thick, mostly 2 mm. thick 
or less. 

Style branches 2; achenes lenticular, black to reddish brown. 

Sheaths firm at the apex. Achene black E. caribaea 

Sheaths membranous at the apex. 

Mature achene black E. maculosa 

Mature achene olivaceous. 

Spikelets of the same diameter as the culms; bristles shorter than the 
achene, glistening white E. Sellowiana 

Spikelets much thicker than the culms; bristles longer than or slightly 
shorter than the achene E. Schaffneri 

Style branches 3; achene trigonous. 
Achene longitudinally costate. 

Spikelets 3-6-flowered E. nervala 

Spikelets 6-12-flowered E. radicans 

Achene not longitudinally costate. 
Achenes conspicuously cancellate, i.e., deeply pitted. 

Achene 1-1.3 mm. long E. retroflexa 

Achene 0.5-0.8 mm. long E. minutissima 

Achenes smooth or reticulate, not pitted. 

Perianth bristles absent; achene 0.5 mm. long E. nigrescens 

Perianth bristles present; achene usually 0.7-1.5 mm. long. 

Plants dwarf, the culms all or mostly 3-7 cm. long E. minima 

Plants relatively tall, the culms all or mostly 10-25 cm. tall or more. 

Achene white E. filiculmis 

Achene yellow, olive, or brown. 



146 FIELDIANA: BOTANY, VOLUME 24 

Style base beak-like, subulate to pyramidal, about one-third as 

long as the achene. 
Spikelets ovoid-fusiform, acute; culms slender, rigid, 1-2 mm. 

broad E. rostellata 

Spikelets cylindric, obtuse to subacute; culms terete, spongy, 

2.5-3 mm. broad E. densa 

Style base broad, much shorter. 

Style base conic or pyramidal; achene evidently reticulate. 

E. montevidensis 

Style base mucroniform, its sides almost parallel; achene 
almost smooth, only faintly reticulate . . . . E. Dombeyana 

Eleocharis caribaea (Rottb.) Blake, Rhodora 20: 24. 1918. 
Scirpus caribaeus Rottb. Descr. & Icon. 24. 1773. Scirpus genicu- 
latus L. Sp. PL 48. 1753 (in part). Eleocharis capitata R. Br. Prodr. 
Fl. Nov. Holl. 225. 1810, not Scirpus capitatus L. Tule; Sintule. 

Marshes or wet fields or thickets, often on mud or sand along 
streams, sometimes in mangrove thickets, ascending from sea level 
to about 1800 meters; Pete"n; Alta Verapaz; Baja Verapaz; Izabal; 
Zacapa; Chiquimula; Jalapa; Escuintla; Guatemala; Solola; Huehue- 
tenango. United States to Mexico, British Honduras, and Panama; 
West Indies and South America; Old World tropics. 

Plants densely cespitose, usually perennial, the slender culms firm, mostly 
15-40 cm. tall, striate and sulcate; sheaths stramineous or brown, the apex firm, 
oblique, often attenuate; spikelets subglobose or ovoid, obtuse, many-flowered; 
scales ovate-orbicular, almost cartilaginous to membranous, yellow to pale brown; 
style 2-fid; stamens 2-3; achene 1 mm. long, obovoid, lustrous black, the style base 
spongious, whitened, variable in shape but usually much depressed; bristles 6 or 8, 
coarse, brown, exceeding the achene, rarely absent. 

A decoction of the thick roots is used in Huehuetenango as a 
beverage, with the addition of atol or other substances. 

Eleocharis cellulosa Torr. Ann. Lye. N. Y. 3: 298. 1836. 

Pete"n (Lago de Pete"n); Guatemala (Laguna de Amatitlan), in 
shallow water. Southern United States and Mexico; Bermuda; 
British Honduras (Honey Camp, Lundell 575) ; West Indies. 

A stout perennial with short rhizomes, the culms mostly terete, 30-75 cm. tall; 
upper sheaths purplish, with an elongate mucronate tip; lowest sheaths mem- 
branous and inflated or leaflike; spikelets cylindric, 1.5-4.5 cm. long, obtuse, 
thicker than the culm; scales orbicular or obovate, obtuse, 4 mm. long, rigid, stri- 
ate, stramineous, with a conspicuous brown border and white scarious margins; 
style 3-fid; stamens 3; achene brown, lustrous, elliptic to obovate, lenticular, with 
about 20 rows of quadrangular cells, bearing at the summit a stout spongy beak, 
tipped by the short dark style base; bristles light brown, equaling the achene, 
without teeth. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 147 

Eleocharis densa Benth. PI. Hartw. 27. 1840. Tule. 

Swampy ground along mountain stream, at 2500 meters or less; 
Jalapa (between Miramundo and summit of Montana Miramundo, 
Steyermark 32722). Central Mexico. 

A coarse perennial from a thick creeping rhizome, the culms terete, spongy, 
green, 55-70 cm. tall, 2.5-3 mm. thick, not septate; sheaths thin, purplish at base, 
brownish near tip, truncate at apex; spikelets large, cylindric, many-flowered, sub- 
acute or obtuse, 15-35 mm. long; scales appressed, very numerous, with an opaque, 
pale, broad central area, narrow purplish sides, and scarious apex and margins, 
rounded at apex, 2.8-3 mm. long; style trifid; achene 1.5 mm. long, trigonous, 
obovate, pale greenish- or brownish-yellow, shining, lightly punctate-reticulate; 
style base stramineous to pale brown, flattened, broadly lanceolate, one-third as 
long as the achene body; bristles stramineous to pale yellowish-brown, equaling or 
somewhat shorter than the achene. 

The identification of the Guatemalan material is provisional. 
Dr. Svenson indicated that it probably represents an undescribed 
species resembling the mountain form of E. montana (HBK.) R. & S. 
At present it seems best placed with E. densa, a Mexican species with 
which it agrees most closely in characters of non-septate culms and 
lightly punctate-reticulate achenes, which are much less reticulate 
than in either E. elegans or E. montana. 

Eleocharis Dombeyana Kunth, Enum. PI. 2: 145. 1837. E. 
montana Auct., not (HBK.) R. Br. 

Moist soil in the central and western mountains, 1200-3500 
meters; Chimaltenango; Quiche" (San Miguel Uspantan); Huehuete- 
nango. Mexico; Ecuador to Argentina. 

Perennial, with elongate creeping rhizomes, the culms mostly 5-30 cm. tall, 
slender; spikelets ovate to linear-lanceolate, 8-12 mm. long, acute, many-flowered; 
scales often rather lax, ovate-elliptic, usually acute, brown, with yellowish or green 
costa and hyaline margin; style 3-fid; achene 1-1.3 mm. long, obovate, lustrous, 
yellow or brown, trigonous, smooth or very obscurely reticulate; style base mu- 
croniform, acute, usually with almost parallel sides; bristles 4, brown, exceeding 
or shorter than the achene. 

Eleocharis elegans (HBK.) Roem. & Schult. Syst. Veg. 2: 150. 
1817. Scirpus geniculatus L. Sp. PI. 42. 1753, in part; ed. 2. 71. 1762. 
Scirpus elegans HBK. Nov. Gen. & Sp. 1: 226. 1816. E. geniculata 
Roem. & Schult. Syst. Veg. 2: 150. 1817. Sintul; Tul; Tule; Guir- 
nalda (Coban); Camalote (Coban); Cebolla de rio (Quezaltenango) . 
Figure 27. 

Marshes or wet meadows, in ditches, often in sand along streams, 
abundant in many regions, ranging from near sea level to about 



148 



FIELDIANA: BOTANY, VOLUME 24 



2000 meters; Alta Verapaz; Baja Verapaz; Zacapa; Chiquimula; 
Jutiapa; Santa Rosa; Escuintla; El Progreso; Guatemala; Sacate- 
pe"quez; Solola; El Quiche"; Suchitepe'quez; Retalhuleu; Quezalte- 
nango; San Marcos; Huehuetenango. Mexico and British Honduras 
to Panama; West Indies; South America. 




FIG. 27. Eleocharis elegans. a, Top of culm with spikelet (X 2). b, Base of 
culm with rootstock (X %). c, Scale (X 17). d, Achene with bristles (X 36). 



A coarse perennial, from a thick creeping rhizome, the culms terete, spongy, 
green, 10-150 cm. tall or higher, 3-10 mm. thick, septate; sheaths reddish, trun- 
cate and usually mucronate at the apex; spikelets large, lanceolate to cylindric, 
usually acute; scales very numerous, 2 mm. long, not carinate, obtuse, thin, with 
an opaque brown central area and broad, scarious, pale brown margins; style 2-3- 
fid; achene 1.5 mm. long, obovate, biconvex to slightly trigonous, yellow to brown, 
lustrous, lightly punctate-reticulate; style base dark brown, flattened, lanceolate, 
half as long as the body of the achene; bristles deep brown, almost equaling the 
tubercle. 

Known in El Salvador by the names "tul," "tule," and "sintule." 
In Guatemala, as throughout Central America, the tough spongy 
culms of this species are much used for weaving the petates or esteras 
used as mattresses upon wooden beds, also for other kinds of mats 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 149 

that are spread upon the floor or employed for other purposes. To 
those accustomed to the mattresses of the north, they are hard beds 
indeed, but in Central America they are considered comfortable, at 
least when new. After having been in use for some time and having 
become well flattened, they are replaced by new "soft" ones. The 
stems are much used also for tying bunches of flowers and vegetables 
for market. The plant is abundant in many marshy places, where it 
often forms rather dense colonies of considerable extent. Among 
local species it is easy of recognition because of its thick terete 
spongy culms. 

Eleocharis filiculmis Kunth, Enum. PI. 2: 144. 1837. Scirpus 
sulcatus Roth, Nov. PI. 1821, not Petit Thouars, 1811. E. sulcata 
Nees, Linnaea 9: 294. 1834. E. costaricensis Boeckel. Allgem. Bot. 
Zeitschr. 2: 34. 1896. Tule chiquito. 

Marshes, wet meadows, ditches, willow thickets, and sandy 
stream banks, ascending from sea level to 1900 meters; Alta Verapaz; 
Izabal; Jalapa; Jutiapa; Escuintla; Sacatepe"quez; Chimaltenango; 
Quich^ ; Huehuetenango; Retalhuleu; Suchitepe"quez. Mexico and 
British Honduras to Panama; West Indies; South America. 

Perennial, erect from an ascending rhizome, the culm slender, 15-40 cm. tall, 
compressed and sulcate, 1 mm. wide or less, the sheaths purplish brown or stra- 
mineous, acute, sometimes slightly inflated at the apex; spikelets ovoid-cylindric, 
4-10 mm. long, many-flowered; scales obtuse or emarginate, stramineous or reddish 
brown, with a paler keel and scarious margins; achene 1 mm. long, trigonous, shin- 
ing white, often obscurely reticulate or brown-striolate; style base almost as wide 
as the apex of the achene, irregularly pyramidal, white to light brown; bristles 
white, usually equaling the achene. 

Eleocharis fistulosa (Poir.) Link in Spreng. Jahrb. 3: 78. 1820. 
Scirpus fistulosus Poir. in Lam. Encycl. 6: 749. 1804. Tule; Cebolla 
de agua (Chiquimula) ; Cham (Huehuetenango). 

Marshes, often on lake shores, ascending from sea level to about 
2200 meters; Alta Verapaz; Izabal; Chiquimula; Jalapa; Jutiapa; 
Santa Rosa; Huehuetenango. Texas; southern Mexico; El Salvador; 
Honduras; Costa Rica; Panama; Cuba; South America; Asia and 
Africa. 

A rather stout perennial with thick rhizomes, the culms somewhat cespitose, 
acutely trigonous, 40-60 cm. tall, the sheaths brown, membranous, rather loose, 
pointed at the apex; spikelets 1.5-3.5 cm. long, cylindric, acute; scales 4 mm. long, 
stramineous or grayish, broadly ovate, obtuse or acutish, coriaceous, striate, the 
margins erose; style 3-fid; stamens 3; achene 2-2.4 mm. long, obovate, turgid, 
green or pale brown, rough, with about 20 rows of deeply pitted, quadrangular 



150 FIELDIANA: BOTANY, VOLUME 24 

cells, the apex narrowed to a neck about one-third the width of the achene, broad- 
ening again to form the base of the triangular style; bristles coarse, usually exceed- 
ing the achene, toothed. 

Eleocharis interstincta (Vahl) Roem. & Schult. Syst. Veg. 2: 
149. 1817. Scirpus interstinctus Vahl, Enum. PI. 2: 251. 1806. 

Bogs or marshes, 600 meters or less; Pete"n; Izabal; Jutiapa; 
Huehuetenango. Florida and Texas to Mexico, British Honduras, 
and Costa Rica; West Indies; South America. 

A coarse perennial with thick rhizomes, the culms terete, 40-100 cm. tall, about 
5 mm. thick, conspicuously septate; sheaths membranous, pointed; spikelets cylin- 
dric, 1.5-4 cm. long, many-flowered; scales in several ranks, oblong, often acute, 
striate, stramineous or grayish, with scarious margins; style 2-3-fid; stamens 3; 
achene 2 mm. long, rough, yellow or gray, with prominent transverse rectangular 
cells and a pronounced annular thickening at the summit; style base dark brown; 
bristles 6, longer than the achene, stout, compressed, with coarse teeth. 

Eleocharis maculosa (Vahl) Roem. & Schult. Syst. Veg. 2: 154. 
1817. Scirpus maculosus Vahl, Enum. PI. 2: 247. 1806. 

In ditches or along borders of streams and lakes, 1200-2500 
meters; Alta Verapaz; Jalapa. British Honduras; Honduras; West 
Indies; South America. 

Perennial, usually densely cespitose, with elongate stolons, the culms mostly 
low and 7-35 cm. tall, sometimes taller, striate, rigid, slender; apex of the upper 
sheath conspicuously enlarged and scarious; spikelets 5-12 mm. long, ovoid to 
lanceolate, many-flowered; scales densely imbricate, subobtuse, firm, lustrous, pur- 
plish brown, with scarious margins; style 2-fid; stamens 3; achene obovoid, 1.3- 
1.5 mm. long, narrowed at the base, black and lustrous, the surface minutely 
roughened; style base half as wide as the achene, light brown, with a dilated base 
and a narrow subulate beak; bristles 7-8, reddish brown, unequal, some of them 
usually equaling the achene, with very small retrorse teeth. 

A collection from Coban is noteworthy in having small spikes 
borne at the bases of the culms, according to Dr. Svenson, "a char- 
acter not previously seen by me in this group." 

Eleocharis minima Kunth, Enum. PI. 2: 139. 1837. E. Duran- 
dii Boeckel. Allgem. Bot. Zeitschr. 2: 34. 1896. 

Wet soil, often about pools on plains, sometimes on sulphur- 
impregnated slopes (Volcan de Tecuamburro), 300-2000 meters; 
Pete"n; Jalapa; Jutiapa; Santa Rosa; Suchitepe'quez. Southern 
United States to Mexico; British Honduras; El Salvador; Honduras; 
Costa Rica; West Indies; South America. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 151 

Plants small and cespitose, with fibrous roots, the culms capillary, 3-7 cm. tall, 
quadrangular-sulcate, the sheaths conspicuous, light or dark brown, the apex in- 
flated, obtuse, hyaline; spikelets 2-4 mm. long, ovate, with few or many flowers; 
scales ovate-lanceolate, mostly acute, dark brown, with greenish costa and hyaline 
margin; style 3-fid; achene ovate, 0.7-1 mm. long, acutely trigonous, whitish to 
pale or olivaceous brown, lightly reticulate to minutely striate, narrowed at the 
base and apex, the style base brownish or gray, short-pyramidal; bristles incon- 
spicuous, white, obscurely toothed, shorter than the achene, often greatly 
reduced. 

Eleocharis minutissima Britton, Mem. Torrey Club 16: 60. 
1920. 

Collected at Honey Camp, British Honduras, wet sand near mar- 
gin of water hole in pine ridge, William C. Meyer 111; probably 
extending into Pete"n. Cuba. 

Plants densely cespitose, the culms capillary, 1-3 cm. high, somewhat thick- 
ened at the base; spikelets ovoid, 1-2 mm. long, loosely 3-7-flowered; scales 
spreading, ovate, strongly carinate, green to castaneous, with hyaline margins; 
achene 0.5 mm. long, oblong or obovoid, white to gray, obtusely trigonous, cancel- 
late; style base gray, low, triangular-apiculate, half as wide as the achene; bristles 
none. 

The British Honduras collection was identified by Dr. Svenson. 
Although the species is cited by him (N. Amer. Fl. 18: 537. 1957) as 
known only from the type locality, a re-examination of the British 
Honduras specimen confirms Dr. Svenson's original determination. 

Eleocharis montana (HBK.) Roem. & Schult. Syst. Veg. 2: 
153. 1817. Scirpus montanus Roth, Nov. PI. Sp. 29. 1821. Scirpus 
nodulosus Roth, I.e. Eleocharis nodulosa (Roth) Schult. Mant. 
2: 87. 1824. 

Marshes, ditches, wet fields, and sandbars, 800-2000 meters; Alta 
Verapaz; Baja Verapaz; Jalapa; Jutiapa; Santa Rosa; Chimalte- 
nango; Quiche"; Huehuetenango. Southern United States to Mexico, 
British Honduras, and Panama; West Indies; South America. 

A rather stout perennial with a stout creeping rhizome, the culms 1-2 mm. 
thick, 30-80 cm. tall, terete, septate; sheaths elongate, usually stramineous, often 
purplish red at the base, truncate at the apex and distinctly mucronate; spikelets 
many-flowered, oblong-lanceolate, acuminate or rarely obtuse, 8-15 mm. long; 
scales appressed or slightly spreading, obtuse or acute, scarious throughout, light 
or dark brown; style 2-3-fid; achene 1 mm. long, broadly obovate, biconvex, yellow 
to brown or olivaceous, distinctly pitted-reticulate; style base flattened, brown, 
half as wide as the achene, deltoid, acute; bristles ferruginous, equaling or slightly 
exceeding the achene. 



152 FIELDIANA: BOTANY, VOLUME 24 

Eleocharis montevidensis Kunth, Enum. PL 2: 144. 1837. 
E. arenicola Torr. in Engelm. & Gray, Bost. Journ. Nat. Hist. 5: 
237. 1847. 

Reported from Guatemala by Svenson, on the basis of Heyde & 
Lux 3554, the determination somewhat uncertain. United States 
and Mexico; Brazil to Argentina. 

Perennial with creeping rhizomes, the slender culms erect, mostly 5-45 cm. 
tall, striate; upper sheath deep brown at the base; spikelets ovoid to oblong, ob- 
tuse, 4-13 mm. long, many-flowered; scales ovate, obtuse, brownish or yellowish, 
with hyaline margins; style 3-fid; achene 1 mm. long, obovoid, trigonous, with 
obtuse angles, golden yellow to brown, minutely punctulate to finely reticulate, 
lustrous; style base conic, short, sessile at the apex of the achene or sometimes with 
a slight constriction; bristles 4-6, brown, toothed, equaling or shorter than the 
achene. 

Eleocharis mutata (L.) Roem. & Schult. Syst. Veg. 2: 155. 
1817. Scirpus mutatus L. Amoen. Acad. 5: 391. 1759. 

In swamps or wet places, at 1000 meters or less; Chiquimula 
(Steyermark 31155} ; reported from Alta Verapaz. Mexico to British 
Honduras and Honduras; Nicaragua; Panama; West Indies; tropical 
South America; tropical Africa. 

Plants coarse and stout, from a short caudex, the culms acutely trigonous, 
40-100 cm. tall, the sheaths stramineous or light brown, membranous, pointed at 
the apex, often elongate; spikelets 1.5-5 cm. long, cylindric, usually obtuse; scales 
many-ranked, stramineous, thin, orbicular or obovate, with broad membranous 
sides and erose apex, often subcarinate; style 3-fid; stamens 3; achene 1.7-2.3 mm. 
long, elliptic or obovate, lustrous, rather smooth, yellow to brown, with about 24 
rows of shallow, transversely linear cells, surmounted at the apex by an annular 
thickening which merges gradually into the short style base; bristles 6, irregular, 
equaling the achene, brown, with coarse teeth. 

A specimen from Guatemala, identified by Dr. Svenson as be- 
longing to this species (Fassett 28844, from Dept. Santa Rosa), has 
the achenes definitely constricted below the summit into a beak one- 
half the width of the achene and must be identified, therefore, as 
E. fistulosa. The Steyermark 31155 specimen from Dept. Chiquimula 
has immature achenes, but these are not constricted below the 
summit. 

Eleocharis nervata Svenson, Rhodora 31: 204. 1929. 

Wet meadows or banks, sometimes on mud flats, 1200-3000 
meters; Baja Verapaz; Jalapa; Chimaltenango; El Quiche"; Quezal- 
tenango; San Marcos. Southern Mexico; El Salvador; Costa Rica. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 153 

Plants slender and delicate, with slender elongate rhizomes, the culms soft, 
filiform, 3-10 cm. long, obscurely striate and angulate; upper sheaths appressed, 
membranaceous and marcescent, yellowish, hyaline at the apex; spikelets ovate, 
2-3 mm. long, 3-5-flowered ; scales ovate, acute or obtuse, lax at maturity, cari- 
nate, green or the sides sometimes rufescent; style 3-fid; stamens 3; achene 1.2- 
1.5 mm. long, narrowly obovate, longitudinally costulate; style base conic; bristles 
3, scarcely as long as the achene. 

Eleocharis nigrescens (Nees) Steud. Syn. Cyp. 77. 1855. Scir- 
pidium nigrescens Nees, Linnaea 9 : 293. 1843. E. minutiflora Boeckel. 
Bot. Jahrb. 7: 274. 1886. E. nigrescens var. minutiflora Svenson, 
Rhodora39:226. 1937. 

The species and variety have been collected in British Honduras 
(W. C. Meyer 109, from Honey Camp) and Campeche, and are to 
be expected in Pete'n, if not elsewhere in Guatemala. South Caro- 
lina; Mexico; West Indies; Trinidad; Brazil; Africa; Madagascar; 
Australia. 

A cespitose annual with fibrous roots, or a perennial with rhizomes, the culms 
filiform, erect, 3-7 cm. tall or sometimes even higher, obscurely quadrangular- 
sulcate; sheaths red or greenish, the apex marcescent; spikelets many-flowered, 
ovoid, 2-3 mm. long; scales chestnut-brown with a greenish costa, obtuse to emar- 
ginate, scarcely carinate, lax at maturity; style 3-fid; mature achenes trigonous, 
0.6 mm. long, smooth, light yellowish brown, whitened with prominent costulate 
angles, immature achenes with obscure striolate reticulation; style base brown or 
light gray, pyramidal, one-third as wide as the achene; bristles none. 

Eleocharis plicarhachis (Griseb.) Svenson, Rhodora 31: 158. 
1929. Scirpus plicarhachis Griseb. Cat. PI. Cub. 239. 1866. 

Bogs or wet meadows, 600 meters or less; Pete'n; Alta Verapaz; 
Izabal; Jutiapa. Southern Mexico; Cuba; Panama to Argentina 
and Paraguay. 

Perennial, erect from a spongy rhizome, the culms stiff, 25-60 cm. tall, wiry, 
flexuous, striate and sulcate; sheaths usually rigid, 4-8 cm. long, purplish or stra- 
mineous, oblique at the apex; spikelets 1-2 cm. long, about 25-flowered, narrowly 
cylindric, acute; scales loose, 3.5 mm. long, linear, obtuse, striate, with an evident 
costa; style 2-fid; stamens 3; achene biconvex, 2 mm. long, light brown, orbicular 
to obovate, with about 12 or more longitudinal rows of quadrate cells, narrowed at 
the summit and surmounted by a turgid ringlike elevation from which rises the 
deep brown or black, lanceolate style base; bristles 6, longer than the achene, 
compressed, toothed. 

Eleocharis radicans (Poir.) Kunth, Enum. PI. 2: 142. 1837. 
Scirpus radicans Poir. in Lam. Encycl. 6: 751. 1804. 



154 FIELDIANA: BOTANY, VOLUME 24 

In marshes, wet meadows or fields, moist banks, often along 
streams, especially on sandbars, 300-2000 meters; Alta Verapaz; 
Baja Verapaz; Zacapa; Escuintla; Santa Rosa; Guatemala; Sacate- 
pe'quez; Chimaltenango; Solola; Quezaltenango; Huehuetenango. 
United States to Mexico; West Indies; southern South America; 
Hawaii. 

Plants low and slender, with creeping rhizomes, often forming dense mats, the 
culms 3-8 cm. tall, soft and spongy, striate; sheaths membranous, fugacious; spike- 
lets ovate, acute, 3-4 mm. long, 6-12-flowered; scales ovate-lanceolate, the lower 
obtuse, green, striate, the upper often strongly carinate at the apex; style 3-fid; 
stamens 3; achene narrowly obovate, yellowish, 0.7 mm. long, with elevated longi- 
tudinal ridges and about 30-40 close trabeculae in a longitudinal series; bristles 
usually 4, very slender, white, retrorsely toothed, exceeding the achene. 

Eleocharis retroflexa (Poir.) Urban, Symb. Antill. 2: 165. 1900. 
Scirpus retroflexus Poir. in Lam. Encycl. 6: 753. 1804. E. Chaetaria 
Auct., not Roem. & Schult. (an Old World species). 

Wet fields or stream banks, ranging from sea level to about 2600 
meters; Alta Verapaz; Izabal; Chiquimula; Santa Rosa; Escuintla; 
Guatemala; Suchitepe"quez; Quiche"; Quezaltenango. Alabama; Brit- 
ish Honduras to Panama; West Indies; South America. 

Plants probably annual, cespitose, with fibrous roots, often proliferous, the 
culms filiform, usually recurved, 2-5 cm. long, compressed or quadrangular-sulcate; 
sheaths stramineous to reddish, obtuse, scarious and inflated at the apex; spikelets 
with few or many flowers; scales usually spreading in fruit, green, carinate, obtuse 
to acute, often with chestnut or reddish brown sides; style 3-fid; achene 1-1.2 mm. 
long, trigonous, cancellate, costate, obovoid to urceolate, white or stramineous; 
style base light brown, as wide as the body of the achene and one-third as long, 
pyramidal-acuminate, the angles decurrent on the costae of the achene; bristles 
white, shorter than the achene. 

This species grows abundantly along some of the streams in the 
Pacific lowlands, usually on banks that are wet by spray from the 
current. The plants often form extensive, dense and bright green 
carpets in such situations. 

Eleocharis rostellata Torrey, Fl. N. Y. 2: 347. 1843. 

At edges of swamps, 300 meters; Huehuetenango (Cie"naga de 
Lagartero, below Miramar, Steyermark 51526). British Columbia 
and United States; Mexico; West Indies. Andes of Ecuador and 
Argentina. 

Plants erect from a short vertical rootstock; culms wiry, conspicuously flat- 
tened and sulcate, mostly 5-90 cm. long, sometimes longer, 1-2 mm. thick; sheaths 
rigid, truncate or oblique at the apex; spikelets fusiform, acute, 12-20-flowered, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 155 

8-20 mm. long; scales light brown, rigid, ovate, the uppermost acute; stamens 3; 
style 3-fid; achene obovoid, obtusely triangular or plano-convex, olive, lustrous, 
slightly reticulate, 2-3 mm. long, narrowed into the obtuse, pale green, rostrate 
style base, this about one-third as long as the achene; bristles firm, regularly den- 
tate, light brown, equaling the achene. 

Eleocharis Schaffneri Boeckel. Bot. Jahrb. 7: 274. 1886. 

Wet plains and swamp borders, 850-1800 meters; Jalapa; Juti- 
apa; Quiche"; Huehuetenango. Southern Mexico; Honduras; Nicara- 
gua; Costa Rica. 

Plants small and slender, cespitose, with fibrous roots, the culms light green, 
capillary, setaceous, spreading, 3-5 cm. long, sulcate; apex of the upper sheath 
membranous, scarcely inflated, obtuse; spikelets ovate, acutish, 2-3 mm. long, 
7-15-flowered; scales membranous, green, sometimes with light reddish or bronze 
sides, broadly ovate, obtuse or acute; style 2-fid; stamens 2-3; achene 0.7 mm. long, 
olive-green, the surface with elongate black striations; style base very small, flat- 
tened, gray, slightly apiculate, one-fourth as wide as the achene; bristles 6-7, white, 
somewhat shorter than the achene. 

Eleocharis Sellowiana Kunth, Enum. 2: 149. 1837. ?E. Tuerk- 
heimii Boeckel. Gyp. Nov. 1: 16. 1888 (type from Coban, Alta Vera- 
paz, Tuerckheim 1+30}. 

Swamps or wet meadows, 1300-1500 meters; Alta Verapaz; 
Jalapa. El Salvador; Honduras; Costa Rica; South America. 

Culms numerous, erect, somewhat spongy, 8-15 cm. tall, 1.5-2 mm. thick, 
striate; upper sheath with a hyaline fugacious apex; spikelets ellipsoid, 5-10 mm. 
long, acute, many-flowered; scales appressed, oblong, obtuse, scarcely carinate, 
stramineous, with a narrow brown stripe on each side of the costa; style 2-fid; 
stamens 3; achene 0.8-1 mm. long, broadly obovate, turgid-lenticular, olivaceous, 
shining, minutely black-striate; style base yellowish green, short-conic, acute, 
about one-fourth as wide as the achene; bristles 7-8, white, shorter than the achene. 



FIMBRISTYLIS Vahl 

Reference: H. K. Svenson, Scirpeae, N. Amer. Fl. 18: 550-555. 
1957. 

Annuals or perennials, the culms leafy at or near the base; spikelets umbellate 
or capitate, terete, with several to many flowers, subtended by an involucre of one 
to many leaflike bracts; scales spirally imbricate, usually deciduous, all fertile; 
perianth none; stamens 1-3; style 2-3-cleft, its base enlarged, deciduous at ma- 
turity from the apex of the achene; achene lenticular, biconvex, or 3-angulate. 

A genus of about 125 species, in temperate and tropical regions 
of the whole earth, but chiefly of the Old World tropics. Only the 
following species are known from Central America. 



156 FIELDIANA: BOTANY, VOLUME 24 

Style branches 3; achenes trigonous. 

Spikelets globose or subglobose, rounded at the apex, 2-4 mm. long; scales 

rounded or very obtuse at the apex F. miliacea 

Spikelets linear to linear-oblong, much longer than broad, acute to acuminate, 

4-10 mm. long; scales mucronate. 
Culms slender, 0.5-1 mm. wide at the apex, compressed-triangular; leaves 

very slender, 0.5-2.5 mm. wide F. autumnalis 

Culms more robust, 1.5-3 mm. wide at the apex, strongly compressed; leaves 

2-6 mm. wide F. complanata 

Style branches 2; achenes lenticular. 
Spikelets glomerulate. 

Bracts slender, much exceeding the inflorescence; achenes stramineous, trans- 
versely reticulate; small annuals F. Vahlii 

Bracts not exceeding the inflorescence; achenes nearly black, with superficial 

isodiametric reticulation; perennials 1-4 dm. tall F. spathacea 

Spikelets mostly on elongate rays. 

Achenes longitudinally ribbed; plants usually annual F. dichotoma 

Achenes coarsely cellular or deeply pitted on surface; plants perennial. 

F. spadicea 

Fimbristylis autumnalis (L.) Roem. & Schult. Syst. Veg. 2: 
97. 1817. Scirpus autumnalis L. Mant. PI. 2: 180. 1771. 

Sandy soil of the Pacific plains, or in oak forest, at 1400 meters or 
less; Retalhuleu; San Marcos. United States to British Honduras; 
El Salvador; Nicaragua; Cuba. 

A slender annual, the culms 10-40 cm. tall, 0.5-1 mm. thick at the apex, gla- 
brous or antrorsely scaberulous; leaves tufted at the base of the culms, much 
shorter than the culms, 0.5-2.5 mm. wide, glabrous or antrorsely scaberulous, the 
sheaths greenish or brownish; bracts 3-5 and 1-3 cm. long, usually shorter than 
the umbel; umbel simple or decompound; rays of the umbel 4-8, bearing numerous 
spikelets, these ovoid to narrowly cylindric, sessile or peduncled, terete, many- 
flowered, 4-10 mm. long, 1-2 mm. wide; scales ovate to ovate-lanceolate, thin, 
mucronate, dull brown with a greenish keel, 1.5 mm. long; achene trigonous, obo- 
vate, 0.4-0.5 mm. long, buff to pale yellow, smooth to faintly reticulate. 

Fimbristylis complanata (Retz.) Link, Hort. Berol. 1: 292. 
1827. Scirpus complanatus Retz. Obs. Bot. 5: 14. 1789. 

In marshes or wet soil, 300 meters or less; Izabal; Huehuete- 
nango. Mexico; British Honduras; Honduras; Panama; West In- 
dies; South America; Old World tropics. 

Perennial from a thick hard base, the culms 20-60 cm. tall, 1.5-3 mm. thick 
at the apex, glabrous or scaberulous on the angles; leaves clustered at the base of 
the plant, much shorter than the culms, 2-6 mm. wide, the margins smooth or 
antrorsely scaberulous; bracts 3-5, shorter than the inflorescence, unequal, the 
rays 4-8, branched, unequal; spikelets 4-10 mm. long, 2 mm. wide, ovoid to linear, 
terete, many-flowered; scales 1.5 mm. long, ovate or ovate-lanceolate, thin, mucro- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 157 

nate, brown; achene 0.8-0.9 mm. long, trigonous, obovoid, pale yellow, faintly 
reticulate. 

Fimbristylis dichotoma (L.) Vahl, Enum. PI. 2: 287. 1806. Scir- 
pus dichotomus L. Sp. PL 50. 1753. Scirpus annuus All. Fl. Pedem. 
2: 277. 1785. Scirpus diphyllus Retz. Obs. Bot. 4: 15. 1791. Fim- 
bristylis annua (All.) Roem. & Schult. Syst. Veg. 2: 95. 1817. 
Figure 28. 

A common plant of wet or moist soil, often in waste ground or 
ditches, wet fields, or on sand or gravel bars of stream beds, most 
common at low elevations, but ranging from sea level to about 2000 
meters; Pete"n; Alta Verapaz; Izabal; Zacapa; Chiquimula; Jalapa; 
Jutiapa; Suchitepe"quez; Retalhuleu; San Marcos; Quiche"; Huehue- 
tenango. United States to Panama, and generally distributed in 
West Indies and South America; widely dispersed in Old World 
tropics. 

Plants usually annual, but sometimes enduring for more than a single year, the 
culms usually cespitose, 5-50 cm. tall, glabrous or sparsely pilose; leaves much 
shorter than the culms, 1-2 mm. wide, 5-20 mm. long, acute, with involute mar- 
gins, glabrous or pilose, the sheaths greenish or brownish; bracts 3-5, short, un- 
equal; umbel with 3-5 rays, the inflorescence a simple or compound umbel with 
usually numerous spikelets, the culm rarely bearing a single spikelet; spikelets 
4-10 mm. long, 2-3 mm. thick, ovoid or oblong-ovoid, pedicellate; scales broadly 
ovate, 1-nerved, rounded and apiculate at the apex, reddish brown; achene 0.7- 
1 mm. long, obovate, whitish or pale brown, conspicuously striate vertically. 

Most Central American plants are glabrous or nearly so. A form 
with a single spikelet terminating each culm has been collected at 
Zacapa, and at various other places in Central America, but it is 
not common. 

Fimbristylis miliacea (L.) Vahl, Enum. PI. 2: 287. 1806. Scir- 
pus miliaceus L. Syst. Nat. ed. 10. 868. 1759. 

Wet fields or thickets, sometimes in swamps or marshes or along 
streams, chiefly at low elevations, ascending to about 1000 meters; 
Petn; Izabal; Zacapa; Jutiapa; Santa Rosa; Suchitepe"quez; Retal- 
huleu. United States to Mexico, British Honduras, and Panama, 
southward in South America; West Indies; Old World tropics. 

A glabrous annual, the culms densely cespitose, 10-70 cm. tall, 1 mm. thick at 
the apex; leaves 10-30 cm. long, often equaling the culms, 1-3 mm. wide, thin, 
scaberulous on the margins, the sheaths green or pale brown; bracts none or minute; 
rays of the umbel 3-6, branched; spikelets 2-4 mm. long, subglobose, brown, 20-40- 
flowered; scales 1 mm. long, ovate, thin, rounded at the apex and shortly mucro- 
nate, carinate; achene 0.5 mm. long, obovoid, pale brown, minutely reticulate. 




FIG. 28. Fimbristylis dichotoma. a, Habit (X 1 A). b, Achene (X 25). 
c, Scale (X 17). d, Head of spikelets (X 4). 



158 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 159 

Fimbristylis spadicea (L.) Vahl, Enum. PI. 2: 294. 1806. Scir- 
pus spadiceus L. Sp. PL 51. 1753. Tul fino (Amatitlan) ; Esparrago 
(Puerto de San Jose"; probably a corruption of Esparto). 

In wet soil about streams or swamps, most often in saline flats 
along the seacoast, ascending to about 1200 meters; British Hon- 
duras and Quintana Roo and to be expected in Pete"n; Zacapa; Juti- 
apa; Escuintla; Guatemala; Retalhuleu; San Marcos. United States 
to Mexico and Panama; West Indies; South America. 

Plants perennial, glabrous, the rhizomes short, thick, and hard; culms few or 
numerous, 30-60 cm. tall or more, 1-2 mm. thick at the apex, septate-nodulose, 
covered at the base by black or dark brown sheaths; leaves 20-60 cm. long, 1-3 mm. 
wide, coriaceous, scaberulous on the margins; bracts shorter or longer than the 
inflorescence, few, 10 cm. long or less; umbel with 3-6 rays, these usually branched, 
unequal; spikelets usually numerous, 5-16 mm. long, 3 mm. or more in diameter, 
terete, oblong-ovate or cylindric, pedicellate, 30-50-flowered ; scales 3-4 mm. long, 
ovate, thick-chartaceous, minutely cuspidate, dark brown; achene 1.5-1.8 mm. 
long, lenticular, obovate, brown to black, coarsely cellular, substipitate, truncate 
at the apex. 

Fimbristylis spathacea Roth, Nov. PI. 24. 1821. 

Salt flats along or near the seashore; Izabal. Southern Mexico 
and British Honduras to Panama; West Indies and South America; 
Old World tropics. 

Plants perennial, glabrous, densely cespitose, with very short, thick rhizomes; 
culms 10-50 cm. tall, 1-1.5 mm. thick at the apex, compressed, smooth, rigid; 
leaves very numerous, forming a dense basal cluster, 5-20 cm. long, 1-3 mm. wide, 
coriaceous, the sheaths brown; bracts 3-4, scarcely longer than the inflorescence; 
rays of the umbel numerous, branched, the whole inflorescence small and compact, 
with very numerous spikelets; spikelets ellipsoid to short-cylindric, 3-6 mm. long, 
1.5-2 mm. wide, subcompressed, 12-20-flowered, pedicellate; scales 1.5 mm. long, 
ovate, thin, rounded at the apex, not cuspidate, light to dark brown with hyaline 
margins; achene lenticular, 0.5-0.8 mm. long, broadly obovate, neither apiculate 
nor stipitate, rugulose with minute reticulation, dark brown to blackish. 

One of the typical beach plants of Central America. 

Fimbristylis Vahlii (Lam.) Link, Hort Berol. 1: 287. 1827. 
Scirpus Vahlii ("Vhalii") Lamb. Tab. Encycl. 1: 139. 1791. 

Open pine ridge, 200 meters or less; British Honduras (Swasey 
Branch, Monkey River, Toledo District, Gentle 3871). Southern 
United States; Nicaragua; South America. 

Cespitose annual with capillary culms 1-15 cm. high, or with the spikelets 
often sessile at the base; leaves filiform, nearly equaling the culms, roughened; 
involucral bracts filiform, much exceeding the capitate clusters of 3-8 spikelets; 



160 FIELDIANA: BOTANY, VOLUME 24 

spikelets oblong-cylindric, obtuse, 4-8 mm. long, many-flowered; scales oblong- 
lanceolate, acuminate, green to dull brown, with prominent midrib; achene obo- 
vate, 0.4 mm. long, stramineous, translucent, prominently reticulate with horizon- 
tally elongated cells. 

FUIRENA Rottboell 

Reference: Henry K. Svenson, Scirpeae, N. Amer. Fl. 18: SOS- 
SOY. 1957. 

More or less pubescent perennial plants, erect or nearly so, with short or elon- 
gate rootstocks, the culms trigonous, leafy from the base to the apex, nodose; leaves 
relatively broad, the sheaths usually covering all or the greater part of the culm; 
inflorescence of terminal and axillary, headlike clusters of spikelets; spikelets many- 
flowered, ovoid; scales spirally imbricate, awned, pubescent, the lower 1-2 empty; 
flowers perfect; perianth of 3 sessile or stipitate scales and sometimes of 3 addi- 
tional barbellate bristles, or of 6 bristles; stamens 3; style 3-cleft, not enlarged at 
the base, deciduous; achene trigonous, obovoid or ovoid, smooth, often stipitate. 

About 30 species in warmer regions of both hemispheres. Prob- 
ably no additional species occur in Central America. Three of the 
species listed here are well illustrated by O'Neill (Carnegie Inst. 
Wash. Publ. 522: 297. /. 3. 1940). 

Perianth of 6 slender retrorsely toothed bristles F. incompleta 

Perianth with 3 inner bristles (scales) ovate-thickened above. 
Three outer bristles present, capillary to filiform. 

Scales of the spikelet with short erect mucro 1 mm. long; spikelets erect. 

F. robusta 

Scales of the spikelet with a long filiform awn; spikelets spreading or reflexed. 

F. simplex 

Three outer bristles lacking, only the thickened scales present. 
Perianth scales membranous, with a slender recurved apical awn. 

F. umbellata 
Perianth scales spongy-thickened at the apex; apical bristle usually lacking. 

F. bulbipes 

Fuirena bulbipes Blake, Contr. U. S. Nat. Herb. 24: 2. 1922. 

Ditches or wet fields, 180 meters or less, Izabal (type collected 
along railroad near Cristina, Blake 7578}; Alta Verapaz. Yucatan; 
Chiapas; British Honduras; Cuba. 

Rootstocks slender, creeping; culms bulbous-thickened at the base, 60-90 cm. 
tall, the lower internodes hispid-pilose, the upper ones glabrous; sheaths of the 
leaves hispid-pilose; blades flat, 3.5-15 cm. long, 5-12 mm. wide, about 6-nerved, 
hispidulous above, hispid-pilose beneath and on the margins; panicles elongate, 
the numerous spikelets sessile in heads of 3-8, ovoid or ellipsoid, 5-6 mm. long; 
scales obovate, pale brown, ciliate, about 2 mm. long, 3-nerved and mucronate; 
perianth scales about equaling the achene, stipitate, obovate-oval, papillose-puber- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 161 

ulous and ciliolate, at the tip contracted into a bulbous-thickened appendage, this 
with a short slender apex; achene obovoid-trigonous, pale brown, lustrous, 0.8 mm. 
long. 

O'Neill suggests that because of the great variation in pubescence 
of this plant, it may be a hybrid, perhaps between F. simplex and 
F. umbellata. Svenson believes that this species may be derived 
from F . umbellata. 

Fuirena incompleta Nees in Mart. Fl. Bras. 2, pt. 1: 107. 1842. 

Boggy ground of swamps, at 1350-2200 meters; Alta Verapaz; El 
Progreso; Jalapa; Huehuetenango. Mexico; Honduras; Panama; 
South America. 

Perennial with creeping rootstocks; culms slender, 40-60 cm. tall; leaves nar- 
row, stiff, 5-12 cm. long, 2-5 mm. wide, glabrous to pubescent; inflorescence of 
small dense heads of 4-10 spikelets in a terminal panicle; spikelets oblong or ellip- 
soid, 8-12 mm. long; scales broadly ovate, dull gray- or plumbeous-green, pubes- 
cent, aristate with an awn 1.5-2 mm. long; bristles usually 6, slender, retrorsely 
barbellate, equaling or exceeding the short-beaked achene. 

Fuirena robusta Kunth, Enum. PI. 2: 185. 1837. 

Margins of bogs or lake shores near sea level; Izabal (shores of 
Lago Izabal, Steyermark 39596). Tabasco; Panama; Cuba; South 
America. 

Plants robust, glabrous or nearly so, 1-2 meters tall, bearing numerous leaves, 
often 5 mm. thick or more, spongy; leaves very large and broad, as much as 30 cm. 
long and 2 cm. wide, some of the leaves shorter, scaberulous on the margins, 5-7- 
nerved, acute; spikelets very numerous, forming a short or elongate panicle, brown, 
ovoid or globose-ovoid, acute, commonly 7-8 mm. long; scales appressed, puberu- 
lent, rounded and short-mucronate at the apex; bristles slender, nearly equaling 
the scales, usually smooth ; perianth scales ovate-lanceolate, slender-stipitate, mu- 
cronate, much longer than the fruit; achene ovoid, smooth, whitish or brownish. 

Fuirena simplex Vahl, Enum. PI. 2: 384. 1806. F. zacapana 
Bartlett, Proc. Amer. Acad. 43: 50. 1907 (type from swamps at 
Gualan, Zacapa, Deam 1*28}. Tubux (Pete"n, Maya). 

In bogs or marshes or along streams, chiefly at low elevations 
but ascending to about 1800 meters; Pete"n; Zacapa; Chiquimula; 
Quiche" ; Huehuetenango. United States to Mexico and British 
Honduras; Nicaragua; Cuba. 

Plants erect, with short, ligneous, creeping, occasionally tuber-bearing root- 
stock; culms usually cespitose, angled, 10-80 cm. tall, glabrous; leaves about 6 on 
each culm, 3-25 cm. long, mostly 3-10 mm. wide, glabrous on the upper surface, 
sometimes sparsely pilose or densely hispid beneath, rough on the margins, long- 



162 FIELDIANA: BOTANY, VOLUME 24 

acuminate, the sheaths green, the upper ones glabrous, the lowest sometimes 
sparsely pilose; spikelets few or numerous in capitate terminal or axillary clusters, 
about 2-8 in each head, 6-15 mm. long, ovoid or oblong-ovoid; scales brown, hispid 
with prominent spreading awns; perianth bristles 3, barbellate, usually longer than 
the achene; perianth scales long-stipitate, obtuse or retuse, short-awned; achene 
yellowish-brown, about 1 mm. long, ovoid, stipitate, the long, somewhat thick- 
ened beak glabrous. 

Fuirena umbellata Rottb. Descr. & Icon. 70. pi. 19, f. 3. 1773. 
Coyolillo mozote (fide Aguilar). 

Wet soil of ditches, swamps, or fields, at or little above sea level ; 
Izabal; Santa Rosa; reported, perhaps in error, from Sacatepe"quez 
(Duefias, Salviri). Southern Mexico and British Honduras to Pan- 
ama, and elsewhere in tropical regions of both hemispheres. 

Plants with short, thick, occasionally tuber-bearing rootstocks, the culms 40- 
150 cm. tall, often 4-angled; leaves about 5-8 on each culm, 6-16 cm. long, com- 
monly 5-25 mm. wide, 5-9-nerved, glabrous on the upper surface, scaberulous 
beneath, the sheaths loose, usually glabrous, green; panicles pilose, 1-5, terminal 
and in the upper leaf -axils; spikelets in few or many capitate clusters, oblong to 
linear-oblong, 6-10 mm. long; scales greenish-brown, about 2 mm. long, obovate, 
rounded to emarginate at the apex, the 3 nerves confluent at the apex and excur- 
rent as a spreading or recurved awn, hyaline, strigillose and sometimes hirsute; 
3 outer bristles none; perianth scales nearly sessile, obovate, hyaline, 3-nerved, 
truncate or rounded at the apex and thin, the midrib excurrent as an uncinate 
mucro; achene about 1 mm. long, ovoid- trigonous, brown or white, stipitate. 

HEMIGARPHA Nees & Arnott 

Small annuals, usually only a few centimeters tall, with tufted, erect or spread- 
ing, almost filiform culms and leaves; spikelets small, terete, terminal, capitate or 
solitary, subtended by an involucre of 1-3 leaflike bracts; scales spirally imbricate, 
deciduous, all fertile, the flowers perfect, a single hyaline scale present between the 
flower and the rachis of the spikelet; bristles none; stamen 1; style 2-cleft, decid- 
uous, the base not enlarged; achene oblong, turgid or lenticular. 

About 5 species in temperate and tropical regions of both hemi- 
spheres. A single species is found in Central America. 

Hemicarpha micrantha (Vahl) Pax in Engl. & Prantl, Pflan- 
zenfam. 2, pt. 2: 105. /. 113. 1887. Scirpus micranthus Vahl, Enum. 
PL 2: 254. 1806. H. subsquarrosa Nees in Mart. Fl. Bras. 2, pt. 1: 
61. 1842. 

On sandbars or in wet soil, scarce, ranging from near sea level to 
about 1400 meters; Guatemala; Suchitepe"quez; Solola (Atitlan). 
United States and Mexico to British Honduras and Costa Rica; 
Guadeloupe and Martinique; South America. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 163 

Plants glabrous, the filiform culms usually numerous, 2-20 cm. tall, weak, 
striate; leaves 1-2 on each culm, often reduced to purplish sheaths or sometimes 
with blades 1-3 cm. long; terminal bract erect, appearing like a prolongation of 
the culm, 1-3 cm. long; spikelets 2-4, terete, ovoid, about 3 mm. long, densely 
many-flowered; scales about 1 mm. long, obovate, mucronulate, reddish brown, 
thin; perianth scale only 0.1 mm. long; achene cylindric or obovate-oblong, 0.5 mm. 
long, purplish brown, terete, minutely cellular. 

Apparently scarce in Guatemala, or perhaps persisting for only 
a short time during the rainy season. Only one collection of the 
plant has been made by the authors. 

HYPOLYTRUM L. Richard 

Coarse perennials with leafy stems and thick rootstocks; inflorescence corym- 
bose-paniculate, the bracts unequal, large, leaflike; spikelets usually numerous, 
several-many-flowered, the flowers perfect or monoecious; scales spirally imbricate, 
some of the lowest ones empty, the flowers complanate; hypogynous scales 2, lat- 
eral, complicate; bristles none; stamens 3 or fewer; style continuous with the ovary, 
not thickened at the base, the stigma branches 2-3, filiform; achene obtusely angu- 
lar or compressed, erostrate, smooth or rugulose. 

About 50 species in the tropics of both hemispheres. Only one is 
known from Central America. 

Hypolytrum Schraderianum Nees in Mart. Fl. Bras. 2, pt. 1: 
65. t. 5. 1842. H. nicaraguense Liebm. Dansk. Vid. Selsk. Skr. 2: 
235. 1851. Figure 29. 

Wet forest or in wooded or rather open swamps of Pete"n; the 
north coast, at or but little above sea level; Pete"n (Rio Pasion); Alta 
Verapaz; Izabal. Tabasco and British Honduras to Panama; Brazil. 

Plants monoecious, glabrous, with short rhizomes, the culms 120 cm. high or 
less, 6-12 mm. thick near the base, trigonous, smooth; leaves 4-6 on each culm, 
60-80 cm. long, 2-3.5 cm. wide, flat, coriaceous, conspicuously costate and nerved, 
rough on the margins, the lower sheaths purplish; bracts 1 or 2, like the leaves but 
smaller; inflorescence a single dense corymb 10-15 cm. broad; spikelets 1.5 mm. 
long, spicate or capitate, strongly compressed, consisting of 3 flowers, the scales 
brownish; stamen 1; achene 3 mm. long, ovoid, terete, acuminate-umbonate, at 
the base broadly cuneate, pale gray, red-puncticulate. 

In Central America confined to the Atlantic coastal region. 

KYLLINGA Rottboell 

Annuals or perennials, the culms slender, trigonous, leafy at the base, the 
leaves sometimes reduced to bladeless sheaths; bracts 2 or more, leaflike, subtend- 
ing the dense, simple or compound heads of small spikelets; spikelets numerous, 
deciduous from the axis of the head at maturity, consisting of only 3-4 scales, the 




FIG. 29. Hypolytrum Schraderianum. a, Achene (X 13). b, Upper part of 
flowering shoot with leaves (X Yi)> c, Head of spikelets (X 10). d, Flower 
(X 20). e, Scale (X 28). /, Achene (X 13). 

164 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 165 

1-2 lowest scales empty, the middle one fertile, the upper empty or staminate; 
scales 2-ranked, carinate; perianth none; stamens 1-3; style 2-cleft, deciduous from 
the achene, not thickened at the base; achene lenticular. 

About 45 species, in tropical and temperate regions of both hemi- 
spheres. No others are known from continental Central America. 
By recent monographers of the family this genus has been combined 
with Cyperus. In habit, however, the group is fairly well marked, 
and it is probably just as well to maintain Kyllinga as a distinct 
genus, as has been done by most botanists for more than a century. 

Leaves all reduced to bladeless sheaths, these with scarious margins . . K. peruviana 
Leaves with well-developed blades. 

Plants annual, the culms densely cespitose K. pumila 

Plants perennial, with well-developed rhizomes, the culms not cespitose. 

Culms stout, about 1.5 mm. thick at the apex; leaves acute, much shorter than 

the culms K. pungens 

Culms very slender, about 0.5 mm. thick at the apex; leaves attenuate, usually 
almost or quite equaling the culms K. brevifolia 

Kyllinga brevifolia Rottb. Desc. & Icon. 13. pi. 4, /. 3. 1773. 
Cyperus brevifolius Hassk. Bort. Bogor. 24. 1844. 

In ditches or wet fields or forest, often along streams, ascending 
from sea level to about 2500 meters, most plentiful at lower eleva- 
tions; Pete"n; Izabal; Zacapa; Jutiapa; Guatemala; Sacatepe"quez; 
Chimaltenango; Huehuetenango; San Marcos; doubtless also in other 
departments. Southern United States to Mexico, British Honduras, 
and Panama, south to Argentina; West Indies; also in the Old World. 

Plant perennial, with slender, short or elongate rhizomes; culms 10-50 cm. tall, 
0.5 mm. thick at the apex; leaves 1-4 to a culm, 1-15 cm. long, 1-3 mm. wide, 
scaberulous on the costa and margins, the sheaths reddish or yellowish; bracts 3-4, 
unequal; spike usually 1, sessile, ovoid or subglobose, 3-8 mm. long; spikelets 
numerous, 2-3.5 mm. long, elliptic or oblong-lanceolate; scales about 2 mm. long, 
ovate or ovate-elliptic, membranous, mucronate, the keel scaberulous; achene 
1 mm. long, lenticular, ellipsoid or obovate-oblong, brown, minutely puncticulate. 

Kyllinga peruviana Lam. Encycl. 3: 366. 1789. K. vaginata 
Lam. 111. 1: 148. 1791. Cyperus peruvianus F. N. Williams, Bull. 
Herb. Boiss. II. 7: 90. 1907. 

Brackish marshes along or near the seashore, Izabal. British 
Honduras to Panama and Colombia; West Indies; tropical Africa. 

A glabrous perennial, the rhizomes often greatly elongate and thick; culms 
about 1 cm. apart, 30-60 cm. tall, 1 mm. thick at the apex, striate; leaves all re- 
duced to sheaths, or rarely with a blade 1 cm. or less in length, the sheaths yellowish 
brown, coriaceous, with membranous margins; bracts 2-4, unequal, short; heads 



166 FIELDIANA: BOTANY, VOLUME 24 

globose, 6-10 mm. in diameter, with very numerous spikelets; scales unequal, the 
longer 3-3.5 mm. long, the shorter 2.5 mm. long, acuminate, the keel smooth; 
achene 1 mm. long, obovate, yellow or light brown, neither stipitate nor rostrate. 

A plant confined to saline soil of coasts. 

Kyllinga pumila Michx. Fl. Bor. Amer. 1: 28. 1803. K. caespi- 
tosa Nees in Mart. Fl. Bras. 2: 12. 1842. Cyperus densicaespitosus 
Mattf. & Kuekenth. Pflanzenreich IV. 20, pt. 2: 597. 1936. Figure 30. 

Marshes or bogs, often in moist thickets or forest, in sandy or 
gravelly places along streams, or sometimes in cultivated ground, 
common in many localities, chiefly at low elevations, but ascending 
to about 1800 meters; Pete"n; Alta Verapaz; Izabal; Zacapa; Chiqui- 
mula; El Progreso; Santa Rosa; Escuintla; Sacatepe"quez; Suchite- 
pe"quez; Retalhuleu; Solola; Quezaltenango; San Marcos; Huehuete- 
nango. United States to Mexico, British Honduras, and Panama, 
south to Argentina; Old World. 

Plants annual, the culms densely cespitose, 5-40 cm. tall, 0.5 mm. thick at the 
apex; leaves 2-3 to a culm, about 2 mm. wide, usually shorter than the culm, 
scaberulous on the margins, the sheaths reddish brown; bracts 3-4, unequal, similar 
to the leaves; heads 4-8 mm. long and 3-4 mm. wide, mostly solitary, globose; 
spikelets 1.5-2 mm. long, very numerous, appressed; scales unequal, one 2 mm. 
long, the other 1.5 mm. long, scaberulous on the keel, pale green or brownish; 
achene 1 mm. long, oblong, yellowish brown, puncticulate, short-stipitate. 

Collections of this and of K. brevifolia from Guatemala have been 
referred to K. odorata Vahl. 

Kyllinga pungens Link, Hort. Berol. 1: 326. 1827. 

Reported from Guatemala (Standley, Field Mus. Bot. 8: 243. 
1931), probably from Izabal. A species of seashores, extending to 
Panama, West Indies, and South America; also in Africa and Asia. 

Perennial, with short scaly rhizomes, the culms usually erect and stout, 30- 
80 cm. tall; basal leaves much shorter than the culm, 3-4 mm. wide, acute; bracts 
2-4 and 6 cm. long or less; head solitary, globose or short-cylindric, 1 cm. thick; 
spikelets lanceolate, greenish; inner scale boat-shaped, carinate, 3-nerved on each 
side. 

LIPOCARPHA R. Brown 

Low annuals, the culms slender, cespitose, leafy at the base; spikelets terete, 
many-flowered, in a terminal head or umbel subtended by an involucre of 1 to 
several bracts; scales firm, spirally imbricate, all fertile or a few of the lowest ones 
empty, at length deciduous; flowers perfect, with a small hyaline scale on each 
side; bristles none; stamens 1-2; style 2-3-cleft, deciduous, its base not enlarged; 
achene plano-convex or trigonous. 




FIG. 30. Kyllinga pumila. a, Habit (X 
c, Flower (X 14). 



b, Head of spikelets (X 4). 



167 



168 FIELDIANA: BOTANY, VOLUME 24 

About 14 species in tropical and warm regions of both hemi- 
spheres. One other species occurs in Panama and Mexico. 

Lipocarpha macula ta (Michx.) Torr. Ann. Lye. N. Y. 3: 288. 
1836. Kyllinga maculata Michx. Fl. Bor. Amer. 1: 29. 1803. L. 
sphacelata Kunth, Enum. PI. 2: 267. 1837. 

Wet fields or in ditches, ranging from sea level to about 1000 
meters; Jalapa; Jutiapa; Suchitepe'quez; Retalhuleu; San Marcos. 
Southern United States to Mexico and Panama; West Indies and 
South America. 

Plants glabrous, with numerous slender roots, the culms densely tufted, smooth, 
striate, 10-25 cm. tall; leaves 5-15 cm. long, 1 mm. wide, much shorter than the 
culms, the sheaths purplish brown; bracts of the involucre 2-3, short or elongate; 
spikelets usually 2-6, sessile at the apex of the culm, 5-8 mm. long, ovoid or oblong- 
ovoid, terete, obtuse; glumes about 1.5 mm. long, obcuneate to lanceolate, thin, 
acute, mottled with purplish or brownish; perianth of 2 glumelike hyaline scales 
enfolding and longer than the achene; stamen 1; achene 1 mm. long, oblong- 
oblanceolate, light brown, puncticulate, apiculate at the apex, spongious-torulose 
at the base. 

RHYNCHOSPORA Vahl 

Reference: G. Kuekenthal, Vor. Mon. Rhynchosporidae, Bot. 
Jahrb. 74: 375-509. 1949; 75: 90-126, 127-195. 1950; 75: 274-314. 
1951. 

Plants chiefly perennial and with rootstocks, sometimes annual, the culms 
leafy, trigonous or terete, the leaves linear, flat or involute; spikelets ovoid to 
oblong or fusiform, variously arranged; scales thin, 1-nerved, spirally imbricate, 
usually mucronate, the lowest ones empty, the upper flowers imperfect, the lower 
ones perfect; perianth of 1-20 (usually 6) bristles, wanting in some species; sta- 
mens usually 3; style 2-cleft, bidentate, or rarely entire; achene lenticular or turgid, 
not trigonous, smooth or transversely rugulose, capped by the persistent base of 
the style (tubercle), or sometimes by the whole style. 

About 200 species of general distribution, most numerous in trop- 
ical regions. A few additional species are known in southern Central 
America. 

Spikelets all crowded in a single dense large head, 1 cm. or more in diameter. 

Bracts 10-30 cm. long; leaves 8-15 mm. wide R. cephalotes 

Bracts 4 cm. long or less; leaves 3 mm. wide or narrower. 

Heads of 2-8 spikelets; bracts of inflorescence scalelike, inconspicuous. 

R. curvula 

Heads of many spikelets; at least one of bracts of inflorescence prolonged and 
longer than the head. 

Plants glabrous; bracts not ciliate R, globosa 

Plants pubescent; bracts ciliate R. barbata 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 169 

Spikelets variously arranged but never in a single head, or if subcapitate, the head 
much less than 1 cm. in diameter. 

Spikelets in globose heads R. cyperoides 

Spikelets not in globose heads. 

Branches of the style short or none, much shorter than the undivided portion. 
Inflorescence composed of individual pyramidal panicles, the spikelets 
mostly alternate; rachilla between the flowers conspicuously produced, 
often flexuous. 
Scales fuscous or rufous-brown. 

Culms stout throughout; spikelets mostly 6-12 mm. long, 1-2 (rarely 3) 

flowers of a spikelet achene-bearing; achene 2 mm. long. 

R. aristata 

Culms slender at the apex; spikelets 4-5 mm. long, 3-4 flowers of a 

spikelet achene-bearing; achene 1 mm. long R. vulcani 

Scales whitish-stramineous or brownish. 

Scales brownish; spikelets 2-4 fasciculately clustered; inflorescence 
mostly broader than long, broadly triangular or pyramidal; cau- 
line leaves few, remote; bristles 4-6, equaling or longer than the 

achene R. dives 

Scales whitish-stramineous; spikelets mostly solitary, alternate; in- 
florescence elongated, as long as or longer than broad; cauline 
leaves numerous, approximate; bristles few, minute or evanescent. 
Spikelets 6-7.5 mm. long; beak elongated-pyramidal, mostly longer 

than the achene; achene 1.5 mm. long R. Tuerckheimii 

Spikelets 4-5.5 mm. long; beak broadly pyramidal, about half the 

length of the achene; achene 0.75-1 mm. long. . . .R. polyphylla 

Inflorescence composed of 2-5 corymbs with fasciculate spikelets; rachilla 

between the flowers short, straight. 

Beak deeply sulcate on both sides, scarcely exceeding the subtending 
scale, nearly or quite as broad as the achene; achene finely punc- 

tulate R. corymbosa 

Beak scarcely or not at all sulcate, exserted from the subtending scale, 
much narrower than the achene; achene strongly transversely undu- 
late-rugose R. triflora 

Branches of the style equaling or longer than the undivided portion. 
Bristles present. 

Achene smooth or faintly reticulate R. fascicularis 

Achene transversely undulate-rugose. 

Tubercle deltoid-attenuate, well exceeded by the bristles; achene with 
the transverse rugulosities and fine striae blurred to often nearly 
smooth R. Marisculus 

Tubercle deltoid, shorter than to barely equaling the bristles; achene 

corrugate R. rugosa 

Bristles none. 

Spikelets only 1-1.5 mm. long R. micrantha 

Spikelets 2.5-10 mm. long. 
Achenes reticulate or smooth. 

Culms and leaves pilose with spreading hairs; achenes reticulate. 

R. hirsuta 



170 FIELDIANA: BOTANY, VOLUME 24 

Culms and leaves glabrous or practically so; achenes smooth or 

minutely longitudinally striate-reticulate R. divergens 

Achenes transversely rugulose. 

Achenes 3 or more in each spikelet. 
Bracts, bractlets, and glumes glabrous; style base depressed and 

almost discoid; glumes very acute R. eximia 

Bracts, bractlets, and lower glumes pilose; style base pyramidal; 

glumes obtuse R. robusta 

Achenes 1-2 in each spikelet. 

Scales whitish; spikelets only 2-5 on each culm R. pusilla 

Scales brownish to fuscous; spikelets usually numerous. 

Achene tridentate at the apex R. setacea 

Achene not tridentate. 

Glumes almost black; achene faintly rugulose, the tubercle 
longer than broad R. Kunthii 

Glumes pale brown; achene strongly rugose, the tubercle 
short, broader than long . . . R. tennis 

Rhynchospora arista ta Boeckel. Flora 40: 36. 1857, and Lin- 
naea 37: 639. 1873. 

Wet forests, damp thickets, and wet banks, frequently in cloud 
forests, 1320-3700 meters; Alta Verapaz; Jalapa; El Progreso; Chi- 
maltenango; Solola; Quezaltenango; Huehuetenango. Mexico; Hon- 
duras; Costa Rica; Panama; West Indies; South America. 

Plants tall and stout, glabrous or nearly so, 4-9 dm. tall, with leafy culms and 
short thick rhizomes; leaves flat, 5-22 mm. wide, smooth or slightly scaberulous; 
spikelets very numerous, in large elongate panicles, mostly solitary and pedicellate, 
oblong-lanceolate and acuminate, 6-13 mm. long or even longer, 2-3-flowered; 
scales brownish-rufescent, mucronate-aristate; bristles 4-5, soon deciduous, shorter 
or slightly longer than the achene, this obovate or broadly ovate, biconvex, smooth 
or cancellate, stramineous or dark brown, with an elongate conic-subulate beak. 

This species has sometimes been confused with R. macrochaeta 
Steud., a South American species with closely crowded spikelets on 
obliquely ascending branches of the inflorescence. 

Rhynchospora barbata (Vahl) Kunth, Enum. PI. 2: 290. 1837. 
Schoenus barbatus Vahl, Eclog. Amer. 2: 4. 1798. 

Open pine forest of the Oriente, 800-1500 meters; Chiquimula; 
Jalapa; Jutiapa. British Honduras to Panama; northern South 
America. 

A densely cespitose perennial, forming small tufts, the slender culms naked, 
15-40 cm. tall, glabrous or sometimes pilose above; leaves usually half as long as 
the culms or shorter, 1-2 mm. wide, pilose with spreading hairs; flower head one, 
globose and subechinate, about 0.5-1.5 cm. in diameter, brown, the basal bracts 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 171 

usually equaling or shorter than the heads, sometimes longer; scales glabrous, in- 
durate and lustrous; bristles twice as long as the achene; achene broadly winged. 

Rhynchospora cephalotes (L.) Vahl, Enum. PI. 2: 237. 1806. 
Scirpus cephalotes L. Sp. PI. ed. 2. 76. 1762. Navajuela. 

Savannas or hilly pine forest of the Atlantic lowlands; Pete"n; 
Alta Verapaz; Izabal. Southern Mexico and British Honduras to 
Panama; Jamaica; South America. 

Plants large and coarse, often a meter high or more, with rhizomes, often form- 
ing dense clumps, the culms leafy; leaves flat, 50 cm. long or less, 4-12 mm. wide, 
with very rough margins; inflorescence consisting of a single ovoid head of greenish 
spikelets, 2-4 cm. long, subtended by large leaflike bracts; spikelets about 7 mm. 
long; bristles 6, longer than the achene; achene 1.5 mm. long, reticulate, castane- 
ous, the slender beak longer than the body. 

The coarse sawlike edges of the leaves can cut the flesh easily. 
The plant is particularly plentiful on the hills above Quirigua hospital. 

Rhynchospora corymbosa (L.) Britton, Trans. N. Y. Acad. 
Sci. 11: 85. 1892. Scirpus corymbosus L. Amoen. Acad. 4: 303. 1788. 
R. aurea Vahl, Enum. PI. 2: 229. 1806. 

Marshes or moist thickets or forest, ascending from sea level to 
about 1380 meters; Alta Verapaz; Izabal. Southern Mexico to Pan- 
ama; West Indies; South America; widely dispersed in tropical re- 
gions of both hemispheres. 

Plants tall and coarse, glabrous or nearly so, the culms trigonous, often 1 meter 
high or more, the rhizomes short and thick or none; leaves equaling the culm, 30- 
60 cm. long and 0.8-1.5 cm. wide, with rough margins; bracts of inflorescence folia- 
ceous; inflorescence large, often 40 cm. long, corymbose-paniculate, the branches at 
length divaricate, the spikelets 6-8 mm. long, clustered in fascicles of 2-5, usually 
2-3-flowered, but maturing a single achene; scales fuscous-ferruginous or dark 
brown, ovate, obtuse, mucronate; bristles 6, ferruginous, longer than the achene; 
achene oblong-obovate, 2-3 mm. long, brown or castaneous, shining, the tubercle 
long-conic, sulcate on each side, equaling the body of the achene. 

Rhynchospora curvula Griseb. Fl. Brit. W. Ind. 574. 1864. 
R. scapigera Boeckel. Linnaea 38: 402. 1874. 

Collected in open pine woods, near Jenkins Creek, Toledo Dis- 
trict, British Honduras, and probably extending into the savannas 
of Pete"n and Izabal. West Indies; northern South America. 

Densely cespitose perennial, the culms several or many, 4-15 cm. tall, filiform, 
trigonous, sulcate, glabrous; leaves basal, rigid, recurved, much shorter than the 
culm, 1.5-4 cm. long, 1-2 mm. wide, carinate, glabrous; sheaths brown; inflores- 
cence terminal, of 2-8-congested spikelets; bracts short, scalelike; spikelets sessile, 



172 FIELDIANA: BOTANY, VOLUME 24 

linear-lanceolate, 6-7 mm. long, 2-flowered; scales 6-7, gray-brown, margins white- 
hyaline, the lower ovate, the upper longer, ovate-lanceolate; bristles 5, rigid, fer- 
ruginous, equaling or longer than the achene, plumose-ciliate for two-thirds of their 
length, antrorsely scabrid at the apex; achene obovate-oblong, biconvex, brown or 
blackish, white-margined, minutely punctulate. 

Rhynchospora cyperoides (Swartz) Mart. Denkschr. Akad. 
Wiss. Muenchen 6: 149. 1816-17. Schoenus cyperoides Swartz, 
Prodr. Veg. Ind. Occ. 19. 1788. Tule. 

Wet meadows or savannas or along swamp and lake margins, 
often in oak forest, at 1700 meters or less; Pete"n; Alta Verapaz; 
Izabal; Jalapa; Jutiapa; Huehuetenango. Southern Mexico to Pan- 
ama and South America; West Indies; tropical Africa. 

Plants perennial, glabrous or nearly so, with short thick rhizomes, the slender 
or rather stout culms 30-80 cm. tall; leaves longer or shorter than the culms, 2-5 
mm. wide; spikelets collected in few or numerous globose and somewhat echinate, 
dark brown, openly paniculate heads 8-12 mm. in diameter; spikelets containing 
a single achene, with about 7 scales; bristles 6, slightly shorter than the achene; 
achene obovoid, narrowed at the base, 1 mm. long, brownish, smooth or trans- 
versely rugulose, the beak subulate, ciliate, equaling the achene. 

Rhynchospora divergens Curtis, Amer. Jour. Sci. II. 7: 409. 
1849. 

Collected near Honey Camp, British Honduras (W. C. Meyer 
106), and probably extending into Pete"n. Florida; West Indies. 

Plants glabrous or nearly so, erect, the culms densely cespitose and forming 
small clumps, very slender, 20-45 cm. tall, wiry, the rhizomes short and thick; 
leaves 20 cm. long or less, shorter than the culms, setaceous, canaliculate, scarcely 
2 mm. wide; spikelets corymbose, the corymbs usually dense and often almost 
headlike, 1-2 cm. broad; spikelets 2-3-fasciculate, 2-3 mm. long, dark brown, con- 
taining 1-2 achenes; scales ovate, subobtuse; stamens 1-2; bristles none; achene 
0.5 mm. long, pale or stramineous, obovoid, sessile, smooth, the beak very short 
and bulblike. 

Rhynchospora dives Standl. in Yuncker, Field Mus. Bot. 17: 
315. 1938. R. orizabensis C. B. Clarke ex Britton, Trans. N. Y. 
Acad. Sci. 11: 85. 1892, nomen nudum; Kuekenthal, Bot. Jahrb. 74: 
406-407. 1949, with description. 

Moist banks, generally in barrancos of the Pacific boca costa and 
the lower slopes of volcanoes, 750-1500 meters; Suchitepe'quez; Que- 
zaltenango; San Marcos. 

Usually coarse, stout, but sometimes more slender perennial, glabrous or nearly 
so, the culms 0.65-1 meter high, acutely triquetrous, smooth, leafy, especially near 
base; leaves elongated, 3-9 mm. wide, generally smooth except for the scabrid mid- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 173 

rib and margins; inflorescence corymbose-paniculate, mostly broader than long, 
the axillary panicles on slender divergent peduncles, 1-3 cm. long; branchlets sub- 
tending spikelets capillary; spikelets rather crowded, 2-4-flowered, mostly 2-4- 
fasciculate, linear-lanceolate, 5 mm. long, short-pedicellate; scales pale brown; 
bristles 4-6, equaling or exceeding the achene; achene subovate, biconvex, pale 
brown, lustrous, obscurely to prominently cancellate, the beak conic-subulate, one- 
half to two-thirds of the length of the achene. 

Rhynchospora eximia (Nees) Boeckel. Linnaea 37: 601. 1873. 
Spermodon eximius Nees in Seem. Bot. Voy. Herald 222. 1854. 

In wet spots on brushy plains, 800-1000 meters ; Jutiapa . Mexico 
to Panama; Cuba; South America. 

Plants glabrous, with fibrous roots, the slender or stout culms 7-45 cm. tall, 
often cespitose, leafy; leaves flat, shorter than or often exceeding the culms, 1-3 
mm. wide; spikelets few or numerous, in small open corymbs, 7-10 mm. long, 
3-8-flowered, usually long-pedicellate, dark brown; scales thin, ovate to lanceolate, 
acute, usually with hyaline margins; bristles none; achene yellow-brown, orbicu- 
lar-obovate, 1-1. 25 mm. long, transversely rugulose, with a very short, depressed 
tubercle. 

Rhynchospora fascicularis (Michx.) Vahl, Enum. 2: 234. 1806. 
Schoenus fascicularis Michx. Fl. Bor. Am. 1: 37. 1803. Schoenus 
distans Michx. Fl. Bor. Am. 1: 36. 1803. Rhynchospora distans 
(Michx.) Vahl, Enum. 2: 235. 1806. R. fascicularis var. distans 
(Michx.) Chapm. Fl. So. U. S. 527. 1860. 

Reported by Kuekenthal from Guatemala. Collected in British 
Honduras without definite locality indicated (H. P. Smart 85), and 
in Stann Creek Valley, British Honduras (Kinloch 217). South- 
eastern and southern United States, from Virginia south along the 
coastal plain west to eastern Texas; Tabasco; West Indies; Venezuela 
and Dutch Guiana. 

Plants erect, perennial, cespitose, the culms mostly slender, 0.4-1.3 meters tall; 
leaves curling to erect, flat to involute, shorter than the culms, 1-4 mm. wide; 
corymbs densely fasciculate; bracts several, mostly setaceous, exceeding the fasci- 
cles; spikelets narrowly ovoid, 3-4.5 mm. long, sessile, 2-4-flowered, containing 1-3 
achenes; scales ovate to lanceolate, conspicuously mucronate to aristate, castane- 
ous or dark brown; bristles 5-6, rudimentary and short (in var. fascicularis) to well 
developed and exceeding the achene (in var. distans), erect, antrorsely serrulate; 
achene ovate to elliptic, chestnut- to blackish-brown, smooth, 1.3-1.5 mm. long; 
tubercle deltoid to deltoid-subulate, one-third to one-half of the length of the achene. 

The British Honduras specimens examined have the bristles 
longer than the achene and are therefore to be referred to var. 
distans. However, considerable variation is encountered in the char- 
acter of the relative length of the bristle with respect to the achene. 



174 FIELDIANA: BOTANY, VOLUME 24 

Rhynchospora globosa (HBK.) Roem. & Schult. Syst. Veg. 2: 
89. 1817. Chaetospora globosa HBK. Nov. Gen. & Sp. 1: 230. 1815. 
?R. epiglobosa Clarke, Kew Bull. Add. Ser. 8: 34. 1908 (type from 
British Honduras). 

Common on savannas near Cristina, Izabal, 70 meters; Pet&i. 
Oaxaca to British Honduras; Costa Rica; Panama; Cuba; South 
America. 

Plants glabrous or nearly so, densely cespitose from stout hard rhizomes, the 
culms 30-90 cm. tall, slender but stiff and wiry; leaves basal, 15-50 cm. long, 5 mm. 
wide or less, erect, stiff and rigid, blackish brown and indurate at the base; heads 
terminal, solitary, subglobose, 1-2 cm. long, one of the bracts longer than the head 
and erect, the others shorter; spikelets brownish, the scales indurate and lustrous, 
mucronate; bristles about 6; achene fuscous, with a short beak. 

Rhynchospora hirsuta Vahl, Enum. PI. 2: 231. 1806. Schoenus 
hirsutus Vahl, Eclog. Amer. 1: 6. 1796. 

Collected at All Pines (Schipp 676} and at Belize (Bartlett 11254), 
British Honduras, and to be expected in the savannas of Pete'n and 
Izabal. Reported from Panama; West Indies; northern South 
America. 

Plants erect, usually densely cespitose, with fibrous roots, 10-40 cm. tall, 
usually densely pilose almost throughout, sometimes glabrate, the culms slender, 
leafy; leaves equaling or shorter than the culms, erect, 1-2 mm. wide, flat; corymbs 
2-4 cm. high, few or several and forming a narrow elongate panicle; spikelets 
numerous, 3-4 mm. long, 2-4-fasciculate, brown or castaneous, the scales broadly 
ovate, obtuse, with hyaline margins, the lower ones with a pilose midrib; bristles 
none; achene yellow-brown, 0.5 mm. long, coarsely undulate-rugulose, lustrous, 
whitish or lead-colored. 

Rhynchospora Kunthii Nees, Linnaea 9: 297. 1834, nomen 
nudum; in Mart. Fl. Bras. 2, pt. 1: 147. 1842. 

Wet meadows, 2200-3150 meters; Huehuetenango; San Marcos 
(Volcan de Tacana). Mexico; Brazil and probably elsewhere in 
South America. 

Plants perennial, glabrous or nearly so, from short thick rhizomes, the culms 
slender or rather stout, 15-60 cm. tall, leafy, trigonous; leaves shorter than the 
culms, 2-3 mm. wide, pale, flat, rough-margined; spikelets few or numerous, ar- 
ranged in a few dense and almost headlike corymbs; spikelets sessile or nearly so, 
blackish, 3-4 mm. long, containing usually 2 achenes; bristles shorter than the 
achene, this yellowish, obovoid, transversely rugulose, the beak conic, acute. 

Rhynchospora Marisculus Nees, Linnaea 9: 297. 1834. 

Marshes and bogs, sometimes in pine forest, near sea level and 
ascending to 1500 meters; Izabal; Alta Verapaz; Chiquimula; 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 175 

Huehuetenango. Southern Mexico; Costa Rica; West Indies and 
South America. 

Plants erect, perennial, glabrous or nearly so, a meter high or usually lower, 
with short rhizomes, the culms leafy, slender; leaves 20-40 cm. long, 4-7 mm. wide, 
rigid, pale when dried; corymbs lax, forming a small panicle, the peduncles erect 
or recurved; spikelets brown, 5-7 mm. long, approximate and subfasciculate, con- 
taining 1-2 achenes; scales ovate-lanceolate, acuminate, loosely overlapping; 
bristles 6-7, much longer than the achene; achene short-stipitate, obovoid, trans- 
versely rugulose, 1.4-1.6 mm. long, 1-1.2 mm. wide, the beak triangular-attenuate, 
1.2-1.8 mm. long, about equaling the achene. 

Rhynchospora micrantha Vahl, Enum. PI. 2: 231. 1806. 

Wet meadows or along small streams, sometimes in oak forest, 
400-1400 meters; Zacapa; Chiquimula; Jalapa; Jutiapa; reported 
from Suchitepe"quez. Southern Mexico to British Honduras and 
Panama; widely dispersed in tropical America; western Africa. 

Plants annual, with fibrous roots, very slender, 10-50 cm. high, the culms 
leafy, glabrous, weak; leaves shorter than the culm, 1-3 mm. wide; corymbs sev- 
eral, small, lax, with almost filiform spreading branches; spikelets numerous, 
greenish, 1-2 mm. long, containing a single achene; scales ovate-lanceolate, acute; 
bristles none; achene minute, 0.5-0.75 mm. long, obovoid, yellowish-brown, trans- 
versely rugulose, bearing a depressed tubercle. 

Rhynchospora polyphylla Vahl, Enum. PI. 2: 230. 1806. 
Schoenus polyphyllus Vahl, Eclog. Amer. 2: 5. 1798. 

Wet or moist forest or thickets, often in ravines or along streams, 
sometimes in pine forest, 250-1500 meters; Alta Verapaz; Zacapa; 
Chiquimula; reported from Santa Rosa; Retalhuleu; Huehuetenango. 
Southern Mexico to Panama; West Indies and northern South 
America. 

Plants perennial, glabrous or pubescent, with slender rhizomes, the culms very 
leafy, often a meter long, weak and frequently reclining; leaves mostly equaling 
the culms, often very numerous, 3-8 mm. wide, the margins usually rough, glabrous 
or sparsely pilose; panicles 4-7, small or elongate, narrow, axillary and often very 
dense; spikelets pale, very numerous, 4-5.5 mm. long, the scales ovate-lanceolate, 
pale or stramineous; bristles few, almost obsolete; achene 0.75-1 mm. long, obo- 
void-globose, castaneous, shining, cancellate, the beak broadly pyramidal, about 
one-half the length of the achene. 

Rhynchospora pusilla (Swartz) Griseb. in Getting. Abh. 7: 271. 
1857. Schoenus pusillus Sw. Prodr. 20. 1788; Fl. Ind. Occ. 7, t. 6. 1797. 
Hypolytrum Berterii Spreng. Neue Entd. 1: 241. 1820. Rhyncho- 
spora Berterii (Spreng.) Clarke in Urban, Symb. An till. 2: 119. 
1900. 



176 FIELDIANA: BOTANY, VOLUME 24 

Collected at Honey Camp, British Honduras, and to be expected 
in nearby Pete*n. Greater Antilles. 

A small slender perennial with very slender rootstocks and short stolons; culms 
weak, almost filiform, 2-10 cm. long, glabrous; leaves often exceeding the culms, 
1 mm. wide or less, glabrous, or the sheaths pilose; spikelets borne at the apex of 
the culms, usually 2-6 in a cluster, pale green or whitish, lanceolate, 2-4 mm. long, 
containing 1-2 achenes; scales lanceolate, acute; bristles none; achene rounded- 
obovoid, brown, 1 mm. long, transversely rugulose, the tubercle low-conic. 

Rhynchospora robusta (Kunth) Boeckel. Linnaea 37: 616. 
1873. Dichromena robusta Kunth, Enum. PI. 2: 283. 1837. 

Bogs or wet savannas, at 1350 meters or less; Alta Verapaz; 
Izabal. Southern Mexico to Panama; South America. 

Plants 60-100 cm. tall, with short rhizomes, the culms rather stout, leafy, 
glabrous, smooth; leaves few, 3-5, shorter than the culms, 5-8 mm. wide, rather 
rigid, glabrous or especially the midrib and margins ciliate; corymbs large and 
broad, with elongate rays, the bracts and bractlets pilose, the spikelets clustered 
at the ends of the rays, few or usually numerous, 7-10 mm. long, brown, the scales 
rather thin, densely imbricate, often with scarious margins, tipped by a short or 
elongate mucro, the lower with pubescent mucro; bristles none; achene biconvex, 
suborbicular, 1.5 mm. long, pale or yellowish-brown, lustrous, transversely rugu- 
lose, bearing a rather short, conic, deeply bilobed tubercle about three-fourths the 
length of the achene. 

The collection of Seler 2401 from Coban, Alta Verapaz, was re- 
ported by Loesener as R. velutina var. semihirsuta (Boeckel.) Clarke 
(=R. semihirsuta Boeckel.). This collection, deposited in the Gray 
Herbarium, has been identified by the present authors as R. robusta. 

Rhynchospora rugosa (Vahl) Gale, Contr. Gray Herb. 151: 275. 
1944. Schoenus rugosus Vahl, Eclog. Amer. 2: 5. 1798. R. glauca 
Vahl, Enum. PI. 2: 233. 1806. 

Usually in pine-oak forest, sometimes on open grassy banks, or 
in alpine meadows, ascending from near sea level to about 3100 
meters; Alta Verapaz; Izabal; El Progreso; Jalapa; Huehuetenango. 
Southern Mexico to Panama; widely dispersed in the tropics of both 
hemispheres. 

Plants erect, perennial, glabrous or nearly so, the culms slender or rather stout, 
20-80 cm. tall, the rhizomes short and stout; leaves 50 cm. long or less, usually 
shorter than the culms, 1.5-3 mm. wide; corymbs few, distant, lax, mostly 1-3 cm. 
broad; spikelets ovoid, 3 mm. long, brown or dark brown, containing 1-2 achenes; 
scales ovate, acuminate; bristles about 6, upwardly barbellate, as long as the 
achene; achene castaneous, obovoid, 1.5 mm. long, transversely rugulose, the 
tubercle conic, acute, one-third to one-half as long as the achene. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 177 

Rhynchospora setacea (Berg) Boeckel. Vid. Medd. Kjoeben- 
havn 1869: 159. 1870. Schoenus setaceus Berg, Act. Helv. 7: 130. 
pi 9. 1772. 

Wet savannas or wet slopes in pine forest, 1500 meters or less; 
Alta Verapaz; Izabal; Chiquimula. Panama; West Indies and 
South America. 

Plants annual or perennial, usually cespitose, with fibrous roots, glabrous, the 
culms erect, slender, 5-50 cm. tall; leaves mostly shorter than the culms, 1-1.5 mm. 
wide, stiff; corymbs several, small, distant; spikelets 2-3-fasciculate, brown or 
brownish, narrowly lanceolate, 4-5 mm. long, containing 1-2 achenes; scales ovate- 
lanceolate, acute or mucronate; bristles none; achene obovoid, 1-1.5 mm. long, 
biconvex, 3-dentate at the apex, transversely rugulose, obconic-stipitate, the tuber- 
cle conic, forming the central one of the three apical teeth. 

Rhynchospora tenuis Link, Jahrb. 3: 76. 1820. 

Marshes, wet meadows, or moist banks, most often in pine forest, 
1000-2000 meters; Jalapa; Guatemala; Quiche"; Huehuetenango. 
Mexico to Honduras; West Indies and South America. Figure 31. 

Plants perennial, erect and rather stiff, with short slender rhizomes; culms 
cespitose, very slender, 10-40 cm. tall, glabrous; leaves equaling or shorter than 




FIG. 31. Rhynchospora tenuis. a, Habit (X 
Inflorescence (X 2). d, Two spikelets (X 6). 



b, Achene (X 35). c, 



178 FIELDIANA: BOTANY, VOLUME 24 

the culms, mostly basal, 1-3 mm. wide; corymbs 1-3, axillary and terminal, the 
spikelets rather loosely fasciculate, numerous, pale, 3-4 mm. long, narrowly lanceo- 
late, acute, containing a single achene; scales lanceolate or ovate, acute; bristles 
none; achene obovate or suborbicular, sessile, brownish, transversely rugulose, 
about 1 mm. long; tubercle acute, one-fourth to one-half as long as the achene and 
decurrent on its margins. 

Rhynchospora triflora Vahl, Enum. PI. 2: 231. 1806. 

Wet savannas near Cristina, Izabal, 70 meters, Steyermark 38430. 
West Indies; South America; Old World tropics. 

Plants coarse and stout, 7-9 dm. tall, the culms smooth or scab rid above; 
leaves often greatly elongate, mostly 4-8.5 mm. wide, rather stiff and coriaceous, 
rough-margined; spikelets slender-fusiform, forming diffuse panicles, mostly 3-7- 
fasciculate at the tips of long naked branches 6-8 mm. long, subsessile or shortly 
pedicellate, 2-4-flowered; scales ovate, mucronate, castaneous or ferruginous; 
bristles 6, longer than the achene, this obovate or broadry elliptic, biconvex, 3 mm. 
long, bearing a subulate tubercle much narrower and somewhat longer than its 
length. 

Plants pertaining to this species have sometimes been identified 
as R. inundata Fern. 

Rhynchospora Tuerckheimii C. B. Clarke, Trans. N. Y. Acad. 
Sci. 11: 85. 1892, nomen nudum; Kuekenthal, Bot. Jahrb. 74: 403. 
1949, with description. Paja blanca; Lechuga; Samracdn (Coban, 
Quecchi) . 

Moist forests, wet brushy banks, frequently in cloud forests, 900- 
2000 meters; Chiquimula; Alta Verapaz; Chimaltenango; Quiche"; 
Huehuetenango; San Marcos. Honduras; Costa Rica. 

Culms very leafy, erect to reclining or subscandent, 1-2.2 meters tall, con- 
spicuously triangular, smooth; leaves elongated, distichous, 3.5-6 mm. long, the 
denticulate-scabrid margins often somewhat revolute, sheaths suffused with rose- 
red; panicles 3-7, pyramidal, distant, 2-5 cm. long; bracts foliaceous, exceeding the 
panicles; spikelets terete, 6-7.5 mm. long, 4-5-flowered; scales densely imbricate, 
stramineous- whitish; bristles few, whitish, quickly deciduous; achene broadly obo- 
vate, biconvex, 1.5-1.75 mm. long, pale brown, shining, conspicuously cancellate, 
the beak elongated-pyramidal, longer than the achene. 

Near Barillas, in the department of Huehuetenango, this species 
is a common ground cover in forests of Liquidambar and Pinus 
Ayacahuite. It is often used as a thatch material for small dwellings. 

Rhynchospora vulcani Boeckel. Linnaea 37: 638. 1873. R. 
Hoffmanni Boeckel. Linnaea 37: 637. 1873. R. ecuadorensis C. B. 
Clarke in Engl. Bot. Jahrb. 34, Beibl. 78: 5. 1904. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 179 

Forested slopes, 1500-3000 meters; Quezaltenango; Huehuete- 
nango; San Marcos. Costa Rica, the type from Volcan de Barba; 
Ecuador. 

Plants perennial, erect, glabrous or nearly so, with short, stout, descending 
rhizomes rooting at the nodes, the culms slender, cespitose, rigid, 3-6 dm. tall, 
leafy at the base; leaves 2-4 mm. wide, elongated, strict, often shorter than the 
culms, lower sheaths rufous-brown; panicles mostly 3-4, interrupted, distant, the 
lateral long-pedunculate, the peduncles erect; foliaceous bracts exceeding the pan- 
icles; spikelets numerous, 3-4 mm. long, fuscous or castaneous, the scales ovate- 
lanceolate, acute; bristles 4-6, slightly exceeding the achene; achenes one in each 
spikelet, obovate, biconvex, about 1 mm. long, very pale brownish, shining, can- 
cellate, the subulate-pyramidal pale beak shorter than the achene. 



SCIRPUS L. 

References: Alan A. Beetle, Studies in the genus Scirpus L.: III. 
The American species of the section Lacustres Clarke, Amer. Journ. 
Bot. 28: 691-700. 1941; VI. The section Schoenoplectus Palla, op. 
cit. 30: 395-401. 1943; Scirpus, in N. Am. Fl. 18, 8: 481-504. 1947. 

Plants usually perennial, slender or stout, the culms terete or angulate, leafy 
or the leaves all reduced to sheaths; spikelets terete or somewhat compressed, soli- 
tary, capitate, spicate, or umbellate, subtended by an involucre of one or several 
bracts, or the involucre wanting; scales spirally imbricate, all fertile or the lowest 
sometimes empty; flowers perfect; perianth of 1-6 bristles, sometimes absent; 
stamens 2-3; style 2-3-cleft, not enlarged at the base, deciduous from the achene 
or its base persistent as a subulate tip; achenes trigonous, lenticular, or plano- 
convex. 

About 150 species, distributed in almost all regions of the earth, 
mostly in very wet soil, often in shallow water. Only the following 
species have been found in Central America. 

Spikelet usually one on each culm; culms slender or capillary, 0.5 mm. thick or less. 
Plants aquatic, submersed ; spikelet containing a single achene . . . . S. confervoides 
Plants not aquatic, growing in moist meadows; spikelets containing several 

achenes S. inundatus 

Spikelets several to many on each culm, rarely one; culms mostly 2-10 mm. thick 
or thicker. 

Bracts several, leaflike; perianth bristles none S. cubensis 

Bract one, appearing like a continuation of the culm; perianth bristles present. 
Culms terete; spikelets in numerous heads on each culm. 

Bristles retrorse-scabrous S. validus 

Bristles plumose S. californicus 

Culms acutely 3-angulate; spikelets usually collected in a single head. 

Spikelets acute or at least pointed -at the apex S. americanus 

Spikelets very obtuse at the apex S. chilensis 



180 FIELDIANA: BOTANY, VOLUME 24 

Scirpus americanus Pers. Syn. PI. 1: 68. 1805. S. americanus 
var. longispicatus Britton, Trans. N. Y. Acad. Sci. 11: 78. 1892. 

In mud around springs or small streams, about 200 meters; 
Zacapa (Agua Caliente, between La Fragua and Rio Motagua, 
Steyermark 29221). Widely distributed in North and South Amer- 
ica; in Central America known only from Guatemala; New Zealand; 
Tasmania. 

Plants perennial, from elongate rhizomes or stolons, the culms erect, a meter 
high or often lower, trigonous, smooth, green, with 3 or fewer sheaths at the base, 
these bearing blades 20 cm. long or shorter; involucral bract one, erect, green, 
1.5-12 cm. long, trigonous, scabrous on the angles; spikelets reddish, sessile, gen- 
erally 2-5, sometimes only one, sessile, 5-20 mm. long, 4 mm. broad, acute or 
pointed; scales 4 mm. long, the prominent costa excurrent from the retuse apex; 
style bifid; anther 1.5 mm. long; bristles 4, often tortuous, reddish brown, equaling 
the achene or only half as long, retrorse-barbate; achene dark brown, apiculate, 
plano-convex, smooth, 3 mm. long, 2 mm. broad. 

The single Guatemalan collection has been determined by Beetle. 
It is referable to var. longispicatus. The Maya name (Yucatan) 
is "halal." 

Scirpus calif ornicus (C. A. Mey.) Steud. Norn. Bot. ed. 2. 2: 
538. 1841. Elytrospermum californicum C. A. Mey. Me"m. Acad. 
St. Pe"tersb. Sav. Etrang. 1: 201. pi. 2. 1830. 

In shallow water of lake shores, ranging from sea level to about 
1700 meters; Alta Verapaz; Izabal; Jalapa; Santa Rosa; Guatemala; 
Solola. United States and Mexico; El Salvador; Costa Rica; South 
America; Hawaiian Islands. 

Plants perennial by stout horizontal rhizomes; culms stout, often 2 meters 
high, subterete or obtusely trigonous, naked, spongy; leaves reduced to basal 
sheaths; involucral bract one, short, subulate, erect; umbel compound, appearing 
lateral, with numerous sessile or usually pedicellate spikelets; spikelets ovoid- 
oblong, 6-10 mm. long, acutish, the glumes ovate, brown, mucronate; bristles 
shorter than the achene or equaling it, short-plumose below, barbate above; style 
2-cleft; achene obovate, plano-convex, almost white, short-rostrate, the surface 
cellular-reticulate. 

The plant is particularly abundant at the edge of Lago de Ama- 
titlan, where it forms dense colonies of wide extent. The tough soft 
stems are used for making mats and other similar articles. 

Scirpus chilensis Nees & Mey. Linnaea 9: 293. 1834, nomen; 
Nees in Meyen, Act. Acad. Cur. 10: 93. 1843. S. Olneyi Gray, Bost. 
Journ. Nat. Hist. 5: 238. 1845. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 181 

In marshes or at the edge of water, 1300 meters or lower; Guate- 
mala (Lago de Amatitlan, Standley 61328). United States and Mex- 
ico; British Honduras (Pueblo Nuevo); Costa Rica; West Indies; 
South America. 

A glabrous perennial with rather thick, elongate rhizomes; culms 30-100 cm. 
high, acutely triquetrous, septate-nodulose, smooth, the sheaths brown or purple; 
bract one, erect, 1-3 cm. long; inflorescence a single head of 5-12 spikelets, appear- 
ing lateral; spikelets ovoid or oblong-ovoid, 5-10 mm. long, 2-3 mm. thick, very 
obtuse, 20-30-flowered ; glumes oval to orbicular, brown, emarginate and mucro- 
nate, glabrous, ciliate; perianth bristles usually 6, retro rsely barbellate; style bifid; 
achene 2 mm. long, obovate-cuneate, plano-convex, dark grayish brown, apiculate, 
smooth, lustrous. 

Scirpus confervoides Poir. in Lam. Encycl. 6: 755. 1804. S. sub- 
mersus C. Wright in Sauv. Fl. Cub. 175. 1871. Websteria submersa 
Britton, Bull. Torrey Club 15: 99. 1888. Eleocharis submersa Miq. 
Linnaea 19: 225. 1847. 

Chiquimula, in small waterhole bordering lake near Chiquimula, 
500-1000 meters, Steyermark 30721. Florida; Cuba; South America; 
tropical Africa and Asia. 

Plants aquatic, submersed, glabrous, with slender rhizomes, the culms filiform 
or capillary, usually 5-8 cm. long; leaves 5-8 cm. long, capillary; spikelets terminal, 
solitary, 8 mm. long, oblong-lanceolate, containing a single achene; lowest glumes 
as much as 8 mm. long, membranaceous, pale green; bristles 6-10, longer than the 
achene, retro rsely barbellate; achene obovoid, pale, smooth. 

Apparently a rare plant, known in Central America from a single 
collection, which was determined by Dr. H. K. Svenson. 

Scirpus cubensis Poepp. & Kunth, in Kunth, Enum. PI. 2: 172. 
1837. 

In shallow water of swamps and marshes, sometimes growing in 
floating bogs, as at Lago de Izabal, at or near sea level, and ascending 
(Sacatepe"quez; Huehuetenango) to 1500-1800 meters; Pete"n; Izabal; 
Jutiapa; Sacatepe"quez (fide Hemsley); Huehuetenango. Mexico to 
Panama and Brazil; West Indies; tropical Africa. Figure 32. 

A glabrous perennial with short rhizomes; culms often cespitose, 30-80 cm. tall, 
trigonous, smooth, rigid; leaves 10-50 cm. long, 2-10 mm. wide, flat, sometimes 
rough-margined, the sheaths brownish purple; bracts 4-7, similar to the leaves; 
rays of the umbel 5-8 and 6 cm. long or shorter, unequal; spikelets densely capitate, 
4-8 mm. long, ovoid, obtuse, 6-12-flowered; scales broadly ovate, coriaceous, 
brown, mucronate; perianth none; style bifid; achene 2 mm. long, obovate-oblong, 
plano-convex, pale yellow, lustrous, very minutely cellular. 



182 



FIELDIANA: BOTANY, VOLUME 24 



The Guatemalan material has been referred by Dr. Beetle to var. 
gracilis (Boeckl.) Beetle. 




FIG. 32. Scirpus cubensis var. gracilis. a, Upper part of culm with flowers 
(X Yz). b, Flower and scale (X 10). c, Achene (X 12). d, Head of spikelets 
(X 8). 



Scirpus inundatus (R. Br.) Poir. in Lam. Encycl. Suppl. 5: 103. 
1817. Isolepis inundata R. Br. Prodr. 222. 1810. 

Wet meadows along brooks, 2500 meters; Huehuetenango (east 
of San Mateo Ixtatan, Steyermark 49877 a). Costa Rica; western 
South America; Australia; New Zealand; Malaysia. 

Plants low, perennial, densely cespitose, usually forming very dense clumps or 
mats; culms very slender but stiff, scarcely 0.5 mm. thick, mostly 5-13 cm. high, 
glabrous, pale green, striate-angulate; sheaths bladeless; bract one, erect, equaling 
or shorter than the spikelets; spikelets mostly solitary, sometimes several, or the 
inflorescence often proliferous in age and subumbellate; spikelets ovoid, obtuse or 
subacute, about 4 mm. long; scales ovate, obtuse, dark reddish brown or castane- 
ous, not nerved; style bifid; achene somewhat lenticular, smooth. 

The plant is common in the higher mountains of Costa Rica but 
apparently very rare in Guatemala. 

Scirpus validus Vahl, Enum. PI. 2: 268. 1806. 

Abundant in open swamps, about 1575 meters; Baja Verapaz 
(large swamp below Pan tin, Standley 90991). Widely distributed 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 183 

in North America; Mexico; West Indies; South America; eastern 
Asia; Australia; New Zealand. 

Plants erect from slender rhizomes, the culms sometimes 2 meters high but 
usually lower, sometimes 2 cm. thick, terete; sheaths reddish or pale brown, with 
a stiff narrow blade 5 mm. long or sometimes as much as 10 cm.; bract of the in- 
florescence rigid, 1-7 cm. long; inflorescence usually much branched, the spikelets 
numerous, in clusters of 2-4 on scabrous-marginate pedicels; spikelets ovoid, few- 
many-flowered, 5-10 mm. long, 4-5 mm. broad; scales mucronate, the mucro often 
slightly recurved; bristles brick-red, usually 6, equaling the achene, retrorsely- 
scabrous, tortuous; style 2-fid; anther 2.2 mm. long; achene 2.5 mm. long, 2 mm. 
broad, deep gray when ripe, lenticular. 



SCLERIA Berg 

Reference: Earl C. Core, The American species of Scleria, Brit- 
tonia 2: 1-105. 1936. 

Annuals or perennials, low or tall and coarse, often with rhizomes, the culms 
leafy; flowers monoecious, the staminate and pistillate spikelets separated or borne 
in the same cluster; spikelets small, clustered in terminal or in terminal and axillary 
clusters, sometimes interruptedly spicate; fertile spikelets 1-flowered, the stami- 
nate many-flowered; scales spirally imbricate; perianth none; style 3-cleft, slender 
or sometimes enlarged at the base, deciduous; ovary supported by a disk (hypo- 
gynium), or this wanting; stamens 1-3; achene globose or ovoid, obtuse, crusta- 
ceous or osseous, white or colored, smooth or variously roughened. 

A large genus of about 200 species, half of which are found in 
tropical and temperate America. A few species besides those listed 
below extend into Central America. 

Hypogynium none; inflorescence often an interrupted spike. 
Inflorescence branched. 

Achene smooth S. lithosperma 

Achene obscurely reticulate or verrucose-tuberculate S. micrococca 

Inflorescence simple, spicate. 
Achene reticulate or verrucose. 

Bracts hirsute S. interrupfa 

Bracts glabrous S. verticillata 

Achene smooth or practically so. 

Plants annual S. Lindleyana 

Plants perennial. 

Inflorescence interruptedly glomerate-spicate; scales hirsute. . .S. hirtella 
Inflorescence a single cluster of spikelets at the summit of the culm; 

scales glabrous S. georgiana 

Hypogynium present and conspicuous at the base of the achene. 
Hypogynium with ciliate, fimbriate, or serrate margins. 

Margin of the hypogynium fimbriate or serrate S. latifolia 

Margin of the hypogynium ciliate. 



184 FIELDIANA: BOTANY, VOLUME 24 

Achene large, 3.5-6 mm. long S. macrophylla 

Achene only 1-3 mm. long. 

Achenes 1-2 mm. long; leaves 7-11 mm. wide S. microcarpa 

Achenes 2-3 mm. long; leaves 10-50 cm. wide. 

Style base black, usually persistent S. mitis 

Style base brown, early deciduous S. Eggersiana 

Hypogynium trilobate, the margins entire, not ciliate. 

Inflorescence paniculate, the upper part of the panicle bearing only staminate 
spikelets, the lower part bearing only pistillate spikelets. Achene 
pubescent S. bracteata 

Inflorescence with staminate and pistillate spikelets intermixed throughout. 
Achene smooth. 

Hypogynium with 9 small tubercles S. oligantha 

Hypogynium without tubercles. 
Ligule at the summit of the leaf sheath bearing a conspicuous scarious 

appendage S. secans 

Ligule not appendaged. 

Culms very slender, about 1 mm. thick at the apex S. anceps 

Culms stout, much thicker. 

Lobes of the hypogynium crenulate; achene white, 3-4 mm. long. 

S. setuloso-ciliata 

Lobes of the hypogynium entire; achene white or purplish, 1.5- 
2.5 mm. long. 

Achene white S. pterota 

Achene purple or purplish S. pterota var. melaleuca 

Achene verrucose, reticulate, or papillate. 
Hypogynium not bearing tubercles. 

Achene glabrous S. reticularis 

Achene pilose S. setacea 

Hypogynium bearing 3 tubercles. 

Achenes 1.5 mm. long S. coriacea 

Achene 2-3 mm. long S. ciliata 

Scleria anceps Liebm. Vid. Selsk. Skrivt. V. 2: 257. 1850. 

Moist thickets or pine-oak forest, 700-1600 meters; Jalapa; 
Huehuetenango. Veracruz. 

Perennial, with rhizomes, the culms 30-50 cm. tall, very slender, the angles 
scaberulous or almost smooth; leaves few, 2 mm. wide, scabrous on the margins, 
the sheaths glabrous or nearly so, very narrowly winged; ligule short-ovate, brown, 
pilose; peduncles 1-3, terminal and axillary, few-flowered, the lateral ones capil- 
lary; bracts foliaceous, exceeding the inflorescence; spikelets 3-5 mm. long, the 
scales red-brown or purplish; hypogynium white, tinged with purple, trilobate, the 
lobes ovate, acute, the margin undulate; achene 2-3 mm. long, white, apiculate, 
ovoid, smooth and lustrous, glabrous. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 185 

Scleria bracteata Cav. Icon. 5: 34. pi. 457. 1799. Navajuela. 

Wet thickets or pine forest, ascending to 1450 meters; Alta Vera- 
paz; Izabal. Southern Mexico and British Honduras to Panama, 
southward to Paraguay and Bolivia; West Indies. 

A perennial with thick rhizomes, the culms erect and a meter high or scandent 
and 2-3 meters long or more, very leafy; leaves 15-45 cm. long, 6-18 mm. wide, 
flat, 3-nerved, rough on both surfaces, the margins serrulate-scabrous, tapering 
gradually from the base into a caudate tip, the sheaths purplish, hirsute; ligule 
4 mm. long, ovate, obtuse, stiff, hirsute; inflorescence paniculate, the peduncles 
terminal and axillary, the upper staminate, the lower pistillate, dark purplish 
brown, the rachises scabrous-hirsute; bracts of the pistillate inflorescence 7 cm. 
long or less, leaflike; staminate spikelets several-flowered; pistillate spikelets with 
few or several flowers, the scales purple, acuminate; hypogynium 3-lobate, the 
lobes rounded, with dark purple margin, entire or sometimes dentate; achene white 
or tinged with purple, 2-3 mm. long, subglobose, verrucose or subtuberculate, 
pubescent. 

Called "cutting-grass" and "saw-grass" in British Honduras. 
The plant is particularly abundant on the pine-clad hills around 
Coban, where it often forms extensive and impenetrable tangles, 
frequently in association with Gleichenia. The sharp-edged leaves 
cut the skin and flesh almost like a sharp knife. 

Scleria cilia ta Michx. Fl. Bor. Amer. 2: 167. 1803. 

Reported by O'Neill from El Cayo District, British Honduras, 
and to be expected in Pete"n or Izabal. Southeastern United States; 
West Indies; Honduras. 

Perennial, with a rather stout, nodose rhizome; culms 20-60 cm. tall, erect, 
tufted, rather stout to almost filiform, sharply triquetrous, pilose or almost gla- 
brous; leaves 20-45 cm. long, 2-6 mm. wide, obtuse, ciliate, often much taller than 
the culms, the sheaths pubescent; ligule 2 mm. long or less; inflorescence terminal 
and axillary, few-flowered, the clusters often arising from the lower axils on short 
erect peduncles; bracts elongate, often conspicuously ciliate; staminate spikelets 
many-flowered, the scales purplish, lanceolate, pubescent, ciliate; hypogynium a 
narrow, obtusely 3-angulate border supporting 3 globose brownish tubercles; 
achene 2-3 mm. long, tuberculate, white, globose, fragile, usually mucronate- 
apiculate. 

Scleria coriacea Liebm. Vid. Selsk. Skrivt. V. 2: 259. 1850. 

Zacapa, pine-covered slopes of Sierra de las Minas, 1000-1500 
meters, Steyermark 29684. Also in Guanacaste, Costa Rica. 

Plants perennial, from a thick hard rhizome, the slender culms 15-45 cm. tall, 
triquetrous, rough on the angles; leaves 25-30 cm. long or more, 2-4 mm. wide, 
rigid, erect, scabrous, the sheaths pubescent; ligule ovate, hirsute; inflorescence of 
1-2 few-flowered glomerules, the lateral one remote, on a short erect peduncle; 



186 FIELDIANA: BOTANY, VOLUME 24 

bract foliaceous, erect, appearing like a continuation of the culm; bractlets pubes- 
cent, linear-lanceolate; spikelets 5 mm. long, few-flowered, the scales ovate, fer- 
ruginous, mucronate, ciliate; hypogynium triangular, bearing 3 bilobate whitish 
tubercles; achene transversely rugose-tuberculate, globose, white, 1-2 mm. long. 

Perhaps ultimately to be reduced to synonymy with S. ciliata. 

Scleria Eggersiana Boeckel. Gyp. Nov. 2: 41. 1890. 

Wet fields or thickets at or near sea level ; Izabal ; Pete"n. Panama ; 
Surinam; West Indies. 

Plants rather stout, erect, from thick rhizomes, the culms 1-2 meters tall, 
trigonous, almost smooth; leaves 30 cm. long or more, 1-2.5 cm. wide, scabrous, 
especially on the nerves and margins, coriaceous and rigid, somewhat plicate, the 
sheaths 3-winged, scabrous on the angles, otherwise glabrous; ligule lance-triangu- 
lar, slightly pubescent, rigid, 2.5 mm. long or less; inflorescence paniculate, narrow, 
about 30 cm. long, its branches erect; bracts leaflike, the bractlets setaceous; pistil- 
late scales suborbicular, ferruginous, mucronate; margin of the hypogynium densely 
brown-ciliate, 3-lobate; achene 2.5-3 mm. long, white, subglobose, smooth and 
lustrous, terminated by a conic style base. 

Scleria georgiana Core, Brittonia 1: 243. 1934. 

Collected in British Honduras at several localities, and to be 
expected in adjacent Pete"n. Southeastern United States; Cuba and 
Jamaica. 

Plants with thick horizontal rhizomes, the culms 30-50 cm. tall, slender, wiry, 
glabrous; leaves 12-16 cm. long or more, 1-2 mm. wide, glabrous, linear or filiform 
and resembling the culms; sheaths glabrous, the lower ones without blades; ligule 
none; inflorescence a single terminal fascicle of 2-5 spikelets, the lowest bract 1-8 
cm. long and 1-2 mm. wide, appearing like a continuation of the culm; spikelets 
5 mm. long, the bractlets lanceolate, glabrous; scales red-brown, glabrous, acumi- 
nate; hypogynium none; achene 2 mm. long, white, smooth, but usually with 
longitudinal ridges, dull, 6-porose near the base, ovoid, the base triangular. 

Scleria hirtella Swartz, Prodr. Veg. Ind. Occ. 19. 1788. 

In savannas, wet fields, or bogs, at 1400 meters or less; Alta 
Verapaz; Pete"n; Izabal; Jutiapa. Southern United States and 
Mexico to British Honduras and Panama, southward to Chile and 
Argentina; West Indies; tropical Africa. 

Plants very slender, perennial, with rather stout, aromatic, horizontal rhi- 
zomes, the culms erect, 15-60 cm. tall, glabrous or pilose near the apex, acutely 
trigonous; leaves 4-20 cm. long, 2-5 mm. wide, flat, pubescent, the sheaths hirsute; 
ligule minute or none; inflorescence interruptedly glomerate-spicate, simple, 4-12 
cm. long, with 3-9 sessile, often nutant, remote glomerules; bractlets long- pilose; 
pistillate scales brown, hirtellous; spikelets 4-5 mm. long; hypogynium none; 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 187 

achene 1-2 mm. long, obovoid or subglobose, white, smooth and lustrous, not 
porose at the base or obscurely 9-12-excavate. 

Scleria interrupta L. Rich. Act. Soc. Hist. Nat. Paris 1: 113. 
1792. 

Grassy or rocky hillsides, sometimes in oak forest, 850-1400 
meters; Alta Verapaz; Jalapa; Jutiapa. British Honduras; Hon- 
duras; Cuba; northern South America. 

A slender annual with fibrous roots, the culms 15-50 cm. tall, trigonous, 
sparsely hirsute; leaves 5-20 cm. long, 1-2 mm. wide, pilose, flat, the sheaths 
pilose; ligule minute and with a tuft of hairs, or none; inflorescence glomerate- 
spicate, 5-10 cm. long, with 5-13 glomerules; spikelets 2-4 mm. long, the bractlets 
linear, hirsute; pistillate scales ovate, purplish, mucronate, ciliate on the keel; 
hypogynium none; achene rugose-verrucose or tuberculate, 1-1.5 mm. long, mucro- 
nate, trigonous, 12-porose at the narrowed base. 

Scleria latifolia Swartz, Prodr. Veg. Ind. Occ. 18. 1788. 

Dense wet forest or on wet brushy banks, 500-1600 meters; Alta 
Verapaz j.Chiquimula; Santa Rosa. Extending to Panama, and 
southward to Bolivia and Brazil; West Indies. 

Plants perennial, tall and stout, from thick rhizomes; culms triquetrous, 
sparsely pilose, often 1-1.5 meters tall or even more; leaves 45-60 cm. long, 2-5 
cm. wide, abruptly narrowed at the apex, scabrous on the nerves and margins, 
otherwise glabrous, the sheaths 3- winged; ligule 5-8 mm. long, triangular-ovate, 
rigid, pilose or glabrous; inflorescence large and usually dense, often 15 cm. long 
or more, with rigid branches, the peduncles compressed, scabrous; lowest bract 
foliaceous and 15 cm. long or more, the bractlets setaceous; pistillate scales ovate, 
mucronate, membranaceous, brown, minutely pubescent; staminate spikelets 3-4 
mm. long, lanceolate, short-pedicellate; hypogynium 3-lobate, the lobes deeply 
and irregularly fimbriate; achene depressed-globose, smooth, 3 mm. long, white or 
pale purplish. 

Scleria latifolia var. arundinacea (Kunth) Standl. & Steyerm. 
Field Mus. Bot. 23: 32. 1944. S. arundinacea Kunth, Enum. PI. 2: 
347. 1837. Pajdn bianco. 

Pine forest or in swampy places, sometimes in Liquidambar for- 
est, 2000 meters or lower; Alta Verapaz; Izabal; Huehuetenango. 
British Honduras to Panama, southward to Argentina; Lesser 
Antilles. 

Like the species, but the whole inflorescence dark purple; achenes dark purplish 
or almost black. 

This bears the same relationship to S. latifolia that var. mela- 
leuca does to S. pterota. There are all sorts of intergrading color 



188 FIELDIANA: BOTANY, VOLUME 24 

forms between white and almost black achenes, so that S. arundi- 
nacea perhaps may be regarded as a mere form of S. latifolia. Called 
"navajuela" in Oaxaca. 

Scleria Lindleyana Clarke, Kew Bull. Add. Ser. 8: 56. 1908. 

Moist places in pine forest, 1000-1500 meters; Chiquimula (south- 
east of Conception de las Minas); Jalapa (between Monjas and 
Jalapa). Honduras; French Guiana and Brazil. 

A slender annual with fibrous roots, the culms erect, 10-40 cm. tall, trigonous, 
glabrous or pilose; leaves flat, 20 cm. long or less, 2-4 mm. wide, pilose, the sheaths 
pilose; ligule minute, with a tuft of hairs; inflorescence interruptedly spicate, 5-10 
cm. long, with 5-8 glomerules; spikelets 2-4 mm. long, the bractlets brown, pilose; 
scales dark brown, lanceolate or ovate-lanceolate; hypogynium none; achene 1 mm. 
long, white, smooth and lustrous, globose, mucronate, bearing 4 pores on each side 
of the trigonous base. 

The species has not been recorded previously from North America. 

Scleria lithosperma (L.) Swartz, Prodr. Veg. Ind. Occ. 18. 1788. 
Scirpus lithospermus L. Sp. PI. 51. 1753. 

Savannas, open rocky banks, or wet pine forest, ascending from 
sea level to 1450 meters; Pete"n; Izabal. Florida to Mexico, British 
Honduras, and Panama, southward to Bolivia; Pacific islands and 
tropical Asia. 

Plants perennial, from rather short, nodose rhizomes, erect, the culms very 
slender, triquetrous, glabrous, 30-60 cm. tall; leaves several, 10-20 cm. long, in- 
volute, 1-3 mm. wide, glabrous but scabrous on the margins and keel; sheaths 
pilose or almost glabrous, purplish; ligule short, triangular, pilose; inflorescence 
axillary and terminal, simple or branched, of 1-4 distant interrupted spikes, few- 
flowered; spikelets 3-4 mm. long; bracts filiform, glabrous, the bractlets dark 
brown, glabrous; scales dark brown, ovate-lanceolate; hypogynium none; achene 
2-2.5 mm. long, white, smooth and lustrous, oblong or ovoid-ellipsoid, umbonate, 
the base narrowed, not porose. 

Scleria macrophylla Presl, Rel. Haenk. 1: 200. 1838. S. palu- 
dosa Poepp. & Kunth, in Kunth, Enum. PL 2: 344. 1837. S. macro- 
carpa Salzm. ex Schlecht. Bot. Zeit. 3: 492. 1845, as synonym. 

Swampy thickets, sometimes in Manicaria swamps, at or little 
above sea level; Izabal. Southern Mexico and British Honduras to 
Panama, southward to Brazil and Bolivia. 

Plants erect, perennial from stout rhizomes, coarse and stout, the culms 1-2 
meters tall, sharply triquetrous, smooth, or rough on the angles; leaves 5-7-nerved, 
20-40 cm. long, stiff, attenuate-acuminate, 1-4.5 cm. wide, smooth or somewhat 
roughened on the nerves and margins, the sheaths 3-winged; ligule short, ovate, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 189 

obtuse, rigid; inflorescence paniculate, often much elongate, with erect branches, 
the spikelets densely clustered; bracts foliaceous, the bractlets setaceous, pubes- 
cent; staminate spikelets 4 mm. long, ovoid or oblong-ovoid; staminate scales 
lanceolate, acuminate, minutely pubescent; pistillate scales ovate-orbicular, ciliate 
and minutely pubescent; hypogynium large, undulately 3-lobate, the margin cili- 
ate; achene globose-ellipsoid, white or brownish white, smooth and lustrous, 3.5- 
6 mm. long, umbonate. 

Called "cortadera" in Oaxaca. 

Scleria microcarpa Nees, Linnaea 9: 302. 1834. 

In swamps or marshes, at 1400 meters or less; Alta Verapaz; 
Izabal; Suchitepe"quez. Tabasco to British Honduras and Panama, 
southward to Paraguay; Cuba. 

Plants erect, perennial from rather stout, horizontal rhizomes, the culms 
acutely trigonous, usually about a meter tall, glabrous; leaves 20-40 cm. long, 
7-11 mm. wide, glabrous or somewhat scabrous on the margins, the sheaths 
3-winged, glabrous or nearly so; ligule 1 cm. long or less, lanceolate, rigid, glabrous; 
inflorescence paniculate, the panicles axillary and terminal, erect, very narrow, 
loosely flowered, the lowest bract leaflike, the bractlets linear-subulate, minute; 
staminate spikelets oblong-ovoid, the glumes ovate, obtuse, stramineous; pistillate 
scales ovate-orbicular, stramineous; margins of the hypogynium ciliate; achene 
1-2 mm. long, ellipsoid-ovoid, white, smooth and lustrous, umbonate. 

Scleria micrococca (Liebm.) Steud. Syn. PL Cyp. 179. 1855. 
Hypoporum micrococcum Liebm. Vidensk. Selsk. Skr. V. 2: 256. 1850. 
Scleria areolata Lundell, Amer. Midi. Nat. 29: 471. 1943. 

Known in Guatemala only from the department of Quiche", the 
exact locality unknown (Aguilar 1419). British Honduras (Monkey 
River, Gentle 4047, type of S. areolata) ; Mexico; Costa Rica; Nicara- 
gua; Panama; Cuba; northern South America to Amazonian Brazil. 

Plants annual with fibrous roots; culms erect, slender, 10-50 cm. tall, trigo- 
nous, glabrous or pubescent; leaves 3-27 cm. long, 1-2.5 mm. wide, glabrous or 
pubescent, about equaling or longer than the culm, somewhat involute in drying; 
ligule minute, pubescent; inflorescence 5-15 cm. long, virgately or paniculately 
branched, glomerules interrupted; rachis pilose to glabrate; bracts mostly seta- 
ceous or lower bract foliaceous, bractlets dark brown; spikelets 3 mm. long, the 
scales dark brown, acuminate, the staminate lanceolate, the pistillate ovate-lance- 
olate; hypogynium none; achene 1 mm. long, white, inconspicuously reticulate or 
verrucose, subglobose, apiculate, attenuate, bearing 4-5 pores on each side of the 
trigonous base. 

Dr. Core has studied the type collection of S. areolata and con- 
siders it to be conspecific with S. micrococca. Our examination of 
this collection has confirmed his study. 



190 FIELDIANA: BOTANY, VOLUME 24 

Scleria mitis Berg. Vet. Akad. Handl. Stockh. 26: 145. pi. 5. 
1765. 

Wet thickets or in Manicaria swamps, at or near sea level; Izabal. 
British Honduras to Panama, Bolivia, and Paraguay; West Indies. 

A rather stout perennial from thick woody rhizomes, the culms usually in 
clumps, commonly 1-2 meters tall, triquetrous, smooth or slightly roughened on 
the angles; leaves 60 cm. long or less, 1-2.5 cm. wide, flat, rigid, glabrous but sca- 
brous on the nerves and margins, the sheaths 3-winged; ligule 3 mm. long or less, 
narrow, glabrous; panicles about 30 cm. long, narrow, loosely many-flowered, the 
branches erect; lowest bract leaflike, the very short bractlets setaceous; spikelets 
small, ovoid, the pistillate scales ovate-orbicular, abruptly acuminate; hypogynium 
truncate, densely ciliate with brown or red-brown hairs; achene ellipsoid, 2-3 mm. 
long, smooth, white or sometimes discolored, tipped by the small conic black 
style base. 

Scleria oligantha Michx. Fl. Bor. Amer. 2: 167. 1803. 

Shaded slopes, sometimes in pine forest, 1200-1800 meters; Alta 
Verapaz; Zacapa; Jalapa. Eastern and central United States and 
Mexico; Honduras. 

Plants perennial from an elongate rhizome, the slender culms erect, 30-60 cm. 
tall, acutely trigonous, glabrous or nearly so; leaves 12-25 cm. long, 2-6 mm. wide, 
scabrous on the nerves and margins, somewhat revolute when dry, the sheaths 
narrowly winged, pubescent or glabrous, the lower ones purplish; ligule short, 
ovate, pubescent, scarious-margined; inflorescence of 2-5 dense clusters of spike- 
lets, terminal and axillary, the lateral clusters on long filiform peduncles; bracts 
leaflike, 5 cm. long, ciliate, the bractlets ciliate, pubescent dorsally, linear-lanceo- 
late; scales lance-ovate, purplish, mucronate; hypogynium a narrow, obtusely 
triangular border supporting 8-9 small tubercles; achene 3-4 mm. long, ovoid, 
white and lustrous. 

Scleria pterota Presl, Isis 21: 268. 1826. S. pratensis Lindl. 
ex Nees, Nov. Acta Acad. Leop. Carol. 19, Suppl. 1: 121. 1843. 
Figure 33. 

Moist forest or thickets, often in ditches, chiefly at 500 meters 
or less, but sometimes ascending to about 1300 meters; Alta Vera- 
paz; Santa Rosa; Escuintla; Suchitepe"quez; Retalhuleu; San Marcos. 
Southern Mexico and British Honduras to Panama, southward to 
Argentina; West Indies. 

Plants perennial from a thick nodose elongate rhizome, erect, the culms rather 
slender, mostly 35-60 cm. high but often taller, frequently clustered, sharply 
triquetrous, glabrous, slightly roughened on the angles; leaves 15-40 cm. long, 
5-15 mm. wide, glabrous, scabrous on the margins, flat, the sheaths narrowly 
3-winged; ligule ovate-triangular, 4-10 mm. long, pilose, with cartilaginous mar- 
gin; inflorescence green or yellowish brown, with 3-4 axillary and terminal panicles, 





FIG. 33. Scleria pterota. a, Habit of upper portion of flowering plant ( X ^). 
b, Habit of lower portion of plant (X M). c, Portion of fruiting branch (X 8). 



191 



192 FIELDIANA: BOTANY, VOLUME 24 

the rachises minutely pubescent or almost glabrous, the panicles 5-15 cm. high; 
lowest bract leaflike, equaling the culm, the bractlets filiform; spikelets greenish 
brown or tinged with purple, the staminate scales lanceolate, acuminate; pistillate 
scales ovate-orbicular, abruptly acuminate; hypogynium depressed, 3-lobate, the 
lobes broad, rounded, ciliate or glabrous; achene 1.5-2.5 mm. long, depressed- 
globose or sometimes pointed, smooth, white or yellowish brown, umbonate at 
the apex. 

Scleria pterota var. melaleuca (Reichb.) Uittien, Fl. Surinam 
40. 1934. S. pterota var. melaleuca (Reichb.) Standl. Field Mus. Bot. 
18: 106. 1937. S. melaleuca Reichb. ex Schlecht. & Cham. Linnaea 
6: 29. 1831. 

Wet or moist, mixed or pine forest or in thickets or in moist or 
wet ground generally, chiefly at or near sea level, but ascending to 
about 900 meters; Pete*n; Alta Verapaz; Izabal; Zacapa; Jutiapa; 
Escuintla; Retalhuleu. Southern Mexico and British Honduras to 
Panama, southward to Bolivia and Brazil; West Indies. 

Like the species, but the whole inflorescence often dark purple; achene pur- 
plish to almost black. 

Called "navajuela" in Honduras and "cortadera" or "zacate cor- 
tador" in Oaxaca. Scleria pterota and its variety are the commonest 
representatives of the genus throughout Central America. 

Scleria reticularis Michx. Fl. Bor. Amer. 2: 167. 1803. 

Wet meadows or rocky forest of the Oriente, often in pine-oak 
forest, 800-1800 meters; Chiquimula (Montana Castilla, southeast 
of Quezaltepeque) ; Jalapa (east of Jalapa) ; Jutiapa (between Agua 
Blanca and Amatillo). Eastern and central United States; Mexico; 
Honduras. 

Plants perennial from a short rhizome, erect, the culms very slender, triangu- 
lar, glabrous, 20-50 cm. tall, somewhat scabrous below on the angles; leaves few, 
10-20 cm. long, 1-3 mm. wide, flat, scarcely equaling the culms, the sheaths nar- 
rowly 3-winged, glabrous or nearly so; panicles terminal and axillary, small, the 
lateral ones sessile or on short erect peduncles, few-flowered; bracts leaflike, the 
bractlets linear-setaceous, glabrous; staminate spikelets lanceolate, stramineous; 
pistillate scales ovate-lanceolate, acuminate, stramineous; hypogynium 3-lobate, 
the lobes lanceolate, obtuse, appressed to the base of the achene; achene 1-2 mm. 
long, globose, conspicuously reticulate, dull white, mucronulate. 

The species has not been reported previously from Central 
America. 

Scleria secans (L.) Urban, Symb. Antill. 2: 169. 1900. Schoenus 
secans L. Syst. ed. 2. 865. 1759. Scleria reflexa HBK. Nov. Gen. & 
Sp. 1: 232. 1816. Navajuela; Sec (Quecchi). 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 193 

Moist or wet thickets, at sea level or ascending to about 500 
meters; Pete"n; Alta Verapaz; Izabal; Suchitepe"quez; Huehuete- 
nango. Southern Mexico and British Honduras to Panama and 
Paraguay; West Indies. 

A perennial with rhizomes, the culms suberect and about a meter high or often 
as much as 5 meters long or more and scandent over shrubs and small trees, acutely 
triquetrous, with very rough angles; leaves numerous, long and narrow, 2-5 mm. 
wide, very scabrous on the margins and costa, sometimes also pubescent, the 
sheaths retrorsely scabrous, pubescent toward the apex; ligule usually conspicu- 
ously scarious-appendaged, rounded-ovate, dark brown, glabrous, the margin lac- 
erate; inflorescences terminal and axillary in the upper leaves, the peduncles pilose 
on the angles, the bracts foliaceous, the bractlets setaceous, pilose, the spikelets 
reflexed; pistillate scales ovate, acute, glabrous, dark purple or blackish; hypo- 
gynium flat, suborbicular, almost entire, the margin reflexed and undulate; achene 
2-4 mm. long, globose-ovoid, white, smooth, lustrous, minutely white-pubescent. 

The plant is a great pest where it grows abundantly, for the sharp- 
edged leaves cut the hands and arms painfully, and the tough stems 
form dense tangles. 

Scleria setacea Poir. in Lam. Encycl. 7: 4. 1806. S. hemitaphra 
Steud. Syn. PI. Gyp. 169. 1855. 

In savannas or pine forest, at 1350 meters or less; Izabal; Chiqui- 
mula; Santa Rosa. Eastern and central United States to Mexico, 
British Honduras, Honduras and Panama, southward to Bolivia and 
Brazil; West Indies. 

Plants annual and with fibrous roots or perennial and with a very short rhi- 
zome, the very slender culms trigonous, glabrous, tufted, 15-60 cm. tall, weak and 
sometimes spreading; leaves flat, 1-4 mm. wide, glabrous or sparsely hirsute, some- 
times scabrous on the margins and nerves, the sheaths glabrous or nearly so, 
sometimes narrowly winged; ligule short, rounded and obtuse, pilose; panicles 
terminal and axillary, the lateral ones on long, filiform, often recurved or drooping 
peduncles, loosely flowered; spikelets 2-4 mm. long, the bracts glabrous, linear- 
lanceolate, the bractlets linear-lanceolate or setaceous, glabrous; pistillate scales 
ovate-lanceolate, acuminate, stramineous or tinged with purple; hypogynium 
deeply 3-lobate, the lobes ovate-lanceolate, subacute, appressed; achene 2 mm. 
long, globose-ellipsoid, more or less reticulate, the transverse ridges pilose, dirty 
white, umbonate. 

Called "navajuela" in El Salvador. 

Scleria setuloso-ciliata Boeckel. Flora 65: 30. 1882. 

Wet soil, especially along streams, at 1500 meters or less; Alta 
Verapaz; Chiquimula; Suchitepe"quez; Solola. Veracruz; Honduras; 
Panama; Cuba; Brazil. 



194 FIELDIANA: BOTANY, VOLUME 24 

Plants perennial, the rhizome short or none, the culms about a meter tall, stout, 
erect, triquetrous, retrorsely scabrous on the angles; leaves 15-30 cm. long, 1-2 cm. 
wide, scabrous on the margins and nerves, the sheaths winged; ligule short and 
rounded, with membranaceous margin; panicles terminal and in the upper leaf 
axils, 5-10 cm. long, the bracts leaflike, the bractlets linear, scabrous-ciliate, elon- 
gate; pistillate spikelets stramineous; hypogynium 3-lobate, the lobes appressed, 
rounded, with crenulate margins, white; achene 3-4 mm. long, ovoid, white, smooth 
and lustrous, mucronate. 

Scleria verticillata Muhl. ex Willd. Sp. PI. 4: 317. 1805. 

Moist or dry slopes, usually in pine forest, in the Oriente at 
1000-1500 meters; Zacapa; Chiquimula. Eastern United States and 
Mexico; El Salvador; Honduras; West Indies; Venezuela, Surinam, 
and Brazil. 

Plants very slender, annual, with capillary roots, sometimes fragrant in drying, 
the culms triquetrous, 10-50 cm. tall, glabrous or sometimes minutely pubescent; 
leaves 5-30 cm. long, 1-2 mm. wide, linear or filiform, glabrous, the sheaths usually 
pilose; inflorescence interruptedly glomerate-spicate, simple, of 2-8 erect glom- 
erules; spikelets 2-3 mm. long, castaneous, few-flowered, the bracts glabrous, 6-7 
mm. long, often caudate, the bractlets glabrous; scales oblong-lanceolate, red- 
brown, glabrous, carinate; hypogynium none; achene 1 mm. long, reticulate or 
verrucose, trigonous-globose, white, fragile, mucronate, about 5-porose just above 
the narrowed trigonous base. 

The species has not been recorded previously from Central 
America. 

UNCINIA Persoon 
Reference: Mackenzie, N. Amer. Fl. 18: 6-8. 1931. 

Perennials, the rootstocks short or emitting long slender stolons; culms erect; 
leaves linear, flat or convolute; flowers unisexual, monoecious, in solitary, androgy- 
nous, dense or lax, bracteate or ebracteate spikes, the apical staminate portion 
short; glumes in few to several ranks; perianth none; stamens commonly 3; pistil- 
late flowers each borne in the axis of a glume and surrounded by a perigynium, this 
closed except for a minute orifice; perigynia obscurely trigonous, truncate at the 
orifice; achenes triangular; style slender, usually thickened at the base, included, 
geniculate with the achene; stigmas 3, exserted; rachilla at the side of the achene 
conspicuous, exserted, setiform, uncinate at the apex. 

About 30 species, all except the following confined to Australasia 
and the mountains of South America. One species, U. tennis Poepp., 
extends from South America into the mountains of Costa Rica. 

Uncinia hamata (Swartz) Urban, Symb. Antill. 2: 169. 1900. 
Carex hamata Swartz, Prodr. Veg. Ind. Occ. 18. 1788. Uncinia 
jamaicensis Pers. Syn. PL 2: 534. 1807. U. hamata var. mexicana 
Kuekenth. Pflanzenreich IV. 20: 54. 1909. Sintule; Pajdn. Figure 34. 




FIG. 34. Uncinia hamata. a, Perigynium (X 8). 6, Achene (X 8). c 
Habit (X H). 

195 



196 FIELDIANA: BOTANY, VOLUME 24 

Wet or moist, mixed forest, often in Cupressus forest or even in 
pine forest, 900-3000 meters, chiefly at middle or rather high ele- 
vations; Alta Verapaz; Zacapa (Sierra de las Minas); Jalapa; Saca- 
tepe"quez; Suchitepe"quez ; Chimaltenango; Quezaltenango; San Mar- 
cos. Southern Mexico to Panama, southward to Argentina; Greater 
Antilles. 

Plants densely cespitose, not stoloniferous, the slender culms 25-60 cm. tall, 
obtusely trigonous; leaves 6-15 to each culm, the blades 10-40 cm. long, 2-12 mm. 
wide, rough on the upper surface; spike solitary, ebracteate, linear-cylindric, 6-18 
cm. long, 2.5-3.5 mm. wide; pistillate glumes green, obovate-orbicular to oblong- 
obovate, 4-6 mm. long, obtuse or acutish, closely appressed; perigynia oblanceo- 
late, 4.5-6 mm. long, many-nerved, hispidulous above, the margins ciliate, not 
beaked; rachilla exserted for 3-6 mm., strongly uncinate at the apex. 

The plant seems to be less plentiful in Guatemala than in the 
mountains of Costa Rica. It is a great pest, wherever found, because 
the hooked bristles cling tenaciously to clothing and penetrate the 
skin readily. 

PALMAE. Palm Family 

Reference: B. E. Dahlgren, Index of American palms, Field Mus. 
Bot. 14. 1936. 

Large or small plants, solitary or gregarious, shrublike or treelike, hermaphro- 
dite, monoecious or dioecious, usually polycarpic but sometimes monocarpic, the 
caudex thick or slender, cylindric, emitting at the base simple or branched, epigean 
or hypogean roots, simple or very rarely branched, rarely scandent, usually annu- 
late, unarmed or armed with spines; leaves terminal, or remote along the upper 
part of the caudex, large for the size of the plant, palmately or pinnately cleft, 
rarely bipinnate, or often simple, the segments distinct or confluent; petiole usually 
more or less produced into the blade, dilated at the base into a short or elongate, 
open or closed sheath, the sheath fibrous on the margins or throughout; inflores- 
cence (spadix) long-pedunculate and inserted among the leaves or short-peduncu- 
late and inserted below them, simple or branched, the rachis short or elongate, the 
branches paniculately or flabellately arranged; spadix bearing various spathes, 
these numerous, tubular, and sheathing the peduncle, or more often 2-4, the lower 
ones then incompletely vaginate, the upper one usually much larger, coriaceous or 
ligneous, enclosing the immature spadix, finally splitting ventrally; flowers articu- 
late or continuous with the spadix, sometimes immersed in pits formed of connate 
bracts, solitary or ternate with the central one pistillate, small or rarely large, reg- 
ular and perfect, or unisexual; perianth double, each series trimerous, in the pistil- 
late flower usually accrescent after anthesis; sepals 3, distinct or connate, open or 
imbricate in bud, in the pistillate flower usually broad and imbricate; petals 3, 
distinct or connate, in the staminate flower valvate or rarely imbricate, in the 
pistillate flower convolute-imbricate, rarely valvate at the apex or throughout; 
stamens usually 6, biseriate opposite the sepals and petals, rarely more numerous, 
inserted in the base of the corolla, usually included; staminodia of the pistillate 
flower none or scale-like or subulate or connate to form an annulus or cupule, the 
filaments free or connate, subulate or filiform, rarely complanate; anthers linear, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 197 

oblong, or sagittate, basifixed or dorsifixed, 2-celled, erect or versatile; ovary free, 
in the staminate flower rudimentary or none, 1-celled at the base or perfectly or 
imperfectly 2-3-celled, rarely 4-7-celled, entire, trilobate, or of 3 distinct carpels; 
style none or short, rarely elongate, the stigmas 3, short, erect or recurved, subulate 
or rarely elongate; ovules solitary and erect or sometimes subhorizontal or pendu- 
lous; fruit more or less subtended by the persistent perianth, dry or drupaceous, 
with one or rarely more numerous cells, the exocarp smooth, warty, prickly or 
scaly, the mesocarp usually fibrous, the endocarp membranaceous, crustaceous, 
ligneous, or stone-like; seed free or adherent to the endocarp, the hilum basal or 
lateral, the raphe short or elongate, with numerous simple, branched, or reticulate 
branches; endosperm corneous or cartilaginous, usually oily, solid or hollow, uni- 
form or ruminate. 

The palms constitute a large, almost exclusively tropical, and in 
many respects important group of plants, of distinctive aspect, and 
seldom confused with any other groups except perhaps Cycadaceae 
and Cyclanthaceae. More than any other single group of plants 
they give to the regions in which they grow a peculiar tropical ap- 
pearance. There are, of course, many tropical regions in which no 
palms occur, particularly those of high elevation, although in Guate- 
mala palms do grow in the higher mountains where there is abundant 
moisture and not too great cold, some of the Chamaedoreas ascend- 
ing to about 2700 meters or perhaps even slightly higher. Some of 
the palms, notably the coconut and date, are of great economic im- 
portance in various parts of the earth, but in Guatemala, as in Central 
America generally, they are relatively little used by man. Some do 
have edible fruits and oil-bearing seeds. Others supply edible in- 
florescences or "cabbages," and in the lowlands the larger leaves are 
important thatch material. The so-called palm wine consists of the 
fermented sap of certain large palms. Leaves of many Guatemalan 
palms are used for making hats, mats, and a wide variety of other 
small articles. However, if all Guatemalan palms were destroyed 
over night, the life of the people would be little affected, because for 
most of these purposes other plants could be substituted. 

One of the local industries that consumes a great quantity of palm 
leaves is the manufacture of suyacales (known in Quecchi as mococh), 
sheets made by basting together pinnae of the larger leaves, for use 
by travelers as a protection against rain. In Alta Verapaz these are 
said to be made from the leaves of a palm called "bojmal" or 
"pamac," whose identity is uncertain, but it is certain that the 
leaves of various genera of palms are used for the purpose. Rabinal 
is celebrated for the petate chiapaneco (mats) woven there from palm 
leaves and noted for their thickness, durability, and color designs. 

Taxonomically many of the Central American palms are still im- 
perfectly understood, although about some of the smaller genera 



198 FIELDIANA: BOTANY, VOLUME 24 

there probably is little more to be learned, so far as species represen- 
tation is concerned. The paucity of information regarding some 
palms is caused by the great size of the plants, which makes it im- 
possible to prepare ordinary herbarium specimens, such as have been 
found quite satisfactory in study of most groups of plants. Only in 
the smaller palms, such as Chamaedorea and Reinhardtia do herbar- 
ium specimens give an adequate idea of the appearance of the living 
plant. On this account the Guatemalan palms have been less col- 
lected than most other families. While it is likely that all or most 
of the genera of the region are represented in the following lists, it is 
almost certain that other species await collection and study, espe- 
cially in the large genus Chamaedorea. The palms of Guatemala have 
suffered also from the fact that for some inexplicable reason they 
have been studied almost wholly in Alta Verapaz. While that de- 
partment probably does possess more species of palms than any 
other, palm plants are found throughout the country at low and 
middle elevations, and in some places are much more conspicuous 
than in Alta Verapaz. 

Study of Central American palms has been vastly facilitated in 
recent years by the numerous generic monographs published by 
Burret and the Index of American Palms by Dahlgren. 

Besides the palms formally listed and described here, a good many 
exotic ones are in cultivation in Guatemalan gardens. Since no spe- 
cial attention has been given them by recent collectors, and since 
cultivated palms often are difficult of determination, only a few of 
them can be listed here. 

The following key has been furnished by Dr. H. E. Moore. 
The nomenclature used in the present treatment is in accordance 
with Dr. Moore's latest studies of the various groups concerned. 
He has revised the manuscript and revised the treatment of Chamae- 
dorea, and is currently monographing it. 

Leaves palmate or costapalmate. 

Perianth in one cycle, forming a 6-dentate cup; carpel 1 Thrinax 

Perianth in two cycles; carpels 3 or, in Schippia, 1. 

Carpels 1 or 3, the styles free and recurved ; spathes inflated and sheathing only 
the peduncle, the primary branches of the spadix with deciduous bracts. 

Flowers with a pediceliform base; carpel 1 Schippia 

Flowers without a pediceliform base; carpels 3 Cryosophila 

Carpels 3 and connate by the styles or united; spathes tubular or expanded in 

a blade, sheathing all the primary branches of the spadix. 
Carpels united, forming a depressed-globose fruit with basal stigmatic scar; 
flowers solitary; petiole unarmed Sabal 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 199 

Carpels free, connate by the styles, forming a globose or laterally com- 
pressed fruit, stigmatic remains apical or subapical in fruit. 
Spathes tubular below, expanded in a blade above, equaling the primary 
branches of the inflorescence; fruit pseudostipitate with abortive 

carpels at the apex; perianth glumaceous Washingtonia 

Spathes tubular, sheathing bases of primary branches only, never equal- 
ing the branches; fruit sessile with abortive carpels basal; perianth 
not glumaceous. 
Ovary glabrous; integument not intruded into the endosperm below the 

raphe of the seed; stems cespitose Paurotis 

Ovary pubescent at the very base and young fruit prominently tomen- 

tose; integument intruded into the endosperm below the raphe of 

the seed; stems solitary or more rarely cespitose. 

Flowers in groups of 3 at least at the base of the branchlets; mature 

fruit ellipsoid to subglobose, 20 mm. long or more with blunt 

subapical scar Erythea 

Flowers solitary; fruit ellipsoid, 18 mm. long or less, with apiculate 

stigmatic scar Brahea 

Leaves pinnate or bipinnate. 

Leaves bipinnate, the pinnules shaped like a fishtail Caryota 

Leaves once pinnate. 

Lower pinnae slender, stiff, and spinelike, the leaves induplicate in bud. 

Phoenix 
Lower pinnae not spinelike, the leaves reduplicate in bud. 

Endocarp without pores, the abortive carpels not fused with the fruit. 
Spathes 3 or more. 

Dioecious palms; staminate flowers with valvate or united petals. 

Chamaedorea 
Monoecious palms; flowers borne in groups of 3 or in lines of several 

along the branchlets, the pistillate lowermost. 
Pistillate flowers with imbricate petals; slender palms with 5-10 

flowers in lines Synechanthus 

Pistillate flowers with valvate petals; thickish palms with flowers in 
groups of 3 at least near the base of the branchlets . . Opsiandra 
Spathes 1-2. 

Leaf sheaths forming a conspicuous crownshaft, the spadices infrafoliar. 

Petals of pistillate flowers imbricate; fruit with lateral or subapical 

stigmatic scar Euterpe 

Petals of pistillate flowers valvate; fruit with basal stigmatic star. 

Roystonea 

Leaf sheaths not forming a conspicuous crownshaft, the spadices arising 

among the leaves but sometimes persisting below them. 
Fruit tuberculate with corky points and ridges; pistillate flowers 
larger than staminate and with valvate petals Manicaria 

Fruit not tuberculate; pistillate flowers smaller than staminate and 
with imbricate petals. 

Flowers not immersed in the branches of the spadix; perianth not 
glumaceous; stamens 8-40 Reinhardtia 

Flowers immersed in the branches or axis of the spadix; perianth 
glumaceous; stamens 6. 



200 FIELDIANA: BOTANY, VOLUME 24 

Anthers with parallel locules united to the connective. 

Calyptrogyne 

Anthers with locules separated, divergent, reflexed or pendulous. 
Staminodial tube urceolate with 6 lanceolate radiating lobes; 

ovary with a terminal style Asterogyne 

Staminodial tube cylindric and truncate, entire or more or less 
toothed or laciniate at the apex; ovary with elongate 

lateral or basal style Geonoma 

Endocarp with 3 distinct pores, the abortive carpels fused with the fertile. 
Unarmed palms. 

Fruit very large, to 25 cm. long or more Cocos 

Fruit less than 15 cm. long. 

Petals of staminate flowers rounded and fleshy, linear-subulate or 

narrow; stamens 6; locules of the anthers not twisted . .Scheelea 

Petals of staminate flowers flat; stamens 12-24; locules of the anthers 

separate and divergent, irregularly twisted, inrolled or sinuous. 

Orbignya 
Armed palms. 

Perianth of pistillate flowers divided nearly to the base; staminate 

flowers immersed in the rachilla Acrocomia 

Perianth of pistillate flowers united, cupuliform, tubular or urceolate. 
Staminate flowers immersed in the axis and densely congested at the 

upper part of the axis Astrocaryum 

Staminate flowers not immersed in the axis, mostly scattered among 

the pistillate. 
Terminal pinnae modified into straight or recurved hooks; stems 

vinelike, not self-supporting Desmoncus 

Pinnae never developed into hooks, the rachis covered with seg- 
ments to the apex; stems erect Bactris 



ACROCOMIA Martius 

Reference: L. H. Bailey, Acrocomia preliminary paper, Gentes 
Herb. 4: 420-476. /. 254-298. 1941. 

Tall palms, armed with spines, the trunks solitary, annulate, densely armed 
with long, straight spines; leaves large, terminal, pinnatisect, the segments linear- 
lanceolate, obliquely acuminate, the margins naked, recurved at the base, the costa, 
petiole, and rachis armed with long spines, the sheath short, open; spadices pedun- 
culate, spiny, the branches thick, the pistillate portion flexuous, with remote flowers, 
the staminate portion elongate, cylindraceous; spathes 2, the lower open at the 
apex, the upper ligneous, fusiform, spiny; bracts of the staminate flower confluent, 
those of the pistillate flower membranaceous; staminate flowers numerous, im- 
mersed in the branches, the pistillate few, large, sessile at the bases of the branches; 
staminate sepals minute, scarious, ovate-oblong, obtuse, only slightly imbricate; 
corolla subcylindric, carnose at the base, obtusely trigonous, the segments oblong, 
valvate; stamens 6, the filaments filiform, adnate to the bases of the corolla seg- 
ments, the anthers exserted, linear-oblong, dorsifixed, versatile; pistillate flowers 
much larger than the staminate, ovoid, the perianth little accrescent in fruit, the 
sepals minute, imbricate, the petals longer, coriaceous, free or connate at the base, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 201 

con volute-imbricate; ovary ovoid, 3-celled, the stigmas 3, sessile, subulate-lanceo- 
late, revolute, the ovule horizontal; fruit globose or oblong, glabrous or setulose, 
1-seeded, the stigmas terminal, the pericarp mucilaginous, fibrous within, the endo- 
carp thick, 3-pored at the middle; seed globose or trilobate, affixed to the side of 
the cell, the testa reticulate by the branches of the raphe, the endosperm hard, 
simple, the embryo opposite one of the pores. 

Twenty-five species are recognized by Bailey, ranging from 
Mexico into South America. Two others have been described from 
southern Central America. 

Lower surface of the leaves loosely pubescent A. mexicana 

Lower surface of the leaves almost glabrous, except for a few appressed hairs. 

A. belizensis 

Acrocomia belizensis L. H. Bailey, Gentes Herb. 4: 445. /. 273, 
274- 1941. Suba. 

Occurring at or little above sea level, sometimes growing with 
pines, British Honduras; type from pine ridge, Duck Run, El Cayo 
District, H. H. Bartlett 11561. 

According to Bailey, differing from A. mexicana in having more slender parts, 
the spines much smaller, the pinnae much smaller and not gray or pubescent be- 
neath but only with a few, very small, appressed hairs, the petiole and rachis 
bearing many very small prickles with hairs on the surface, the spadix very slender 
at anthesis, the main axis and rachillas smooth and glossy at maturity, the fruit 
smaller, lustrous, only slightly depressed; pinnae very numerous, the rachis long 
and slender, shallowly sulcate but not flattened, mostly less than 1 cm. broad but 
sometimes 2 cm., gray and dull; costa and nerves of the pinnae prominent, glabrous 
except for a few points and appressed hairs beneath; main spines of the leaf rachis 
9 cm. long or shorter; base of the petiole 3 cm. broad or less, enclosed in a clothlike 
hairy sheath 5 cm. broad or more; main spathe to 20 cm. broad, thick and heavy, 
densely invested with fibrous wool and covered with many flat, very sharp, black 
spines 2-6 cm. long; axis of the spadix glabrous, unarmed, lustrous; flowers small; 
fruit about 3 cm. in diameter, slightly flattened at each end, somewhat lustrous 
but the surface rough. 

It is unknown whether all the Acrocomias of British Honduras 
are referable to A. belizensis, or whether some are A. mexicana. 
Further study will be necessary to decide whether two species of the 
genus are represented in Guatemala and British Honduras, since the 
coyol is widely distributed, and it is difficult to suggest a line at 
which two geographic species might be separated. 

Acrocomia mexicana Karwinsky ex Mart. Hist. Nat. Palm. 3: 
285. pi. 138. 1845. Coyol; Tuc (Maya); Map (Quecchi); Cocoyol; 
Supa (Pet&i). Figure 35. 



202 FIELDIANA: BOTANY, VOLUME 24 

Widely distributed, mostly at 1000 meters or lower, often abun- 
dant in rather open, lowland forest, or on dry open hillsides, often 
abundant on plains, and left when land is cleared; Pete*n; Alta Vera- 
paz; Baja Verapaz; El Progreso; Zacapa; Chiquimula; Jutiapa; Santa 
Rosa; Escuintla; Suchitepe"quez; Solola; Retalhuleu; Quezaltenango; 
San Marcos; Huehuetenango; Quiche". Southern Mexico; probably 
extending into Honduras and El Salvador. 

A large palm, the trunk often 10 meters high, thick and massive, cylindric, 
usually abundantly armed with long sharp dark spines; leaves 4 meters long or 
shorter, often drooping, the dead leaves covering the upper part of the trunk; pin- 
nae close together but unequally spaced ; petiole and rachis hairy and more or less 
setose, armed with flat, brownish black, lustrous spines 11 cm. long or less; pinnae 
at the middle of the leaf 1 meter long or more and about 3 cm. wide, the nerves 
thinly hairy beneath, the rachis spiny throughout; main spathe 70-80 cm. long, 
narrow-pointed at the apex, densely and softly brown-tomentose and often bearing 
slender spines 1 cm. long; spadix about equaling the spathe, the branches glabrous 
or glabrate, the peduncle hairy, setose, and spiny, the branches or rachillae about 
25 cm. long; staminate flowers 8 mm. long at anthesis, the calyx one-fourth or one- 
fifth as long as the deeply obtuse-lobate corolla; anthers exserted; pistillate flowers 
few on each rachilla, about 1 cm. long in anthesis, the styles somewhat exserted; 
corolla tightly involute, curved-striate; fruit depressed-globose, about 3.5 cm. high 
and 4 cm. broad, brownish and somewhat mottled, glabrous, the appressed calyx 
about 1.7 cm. broad. 

This is perhaps the most common native palm of Guatemala, as 
it and its congeners are found through much of Central America. 
It is widely distributed in Guatemala and often conspicuous on the 
landscape, although there are, of course, large areas from which it is 
absent. Frequently it is planted in regions where it is not native, 
for ornament or perhaps sometimes for its fruit. The tree is known 
everywhere by the name "coyol," a word of Nahuatl origin, which 
appears in such place names as Coyoles, El Coyolar, Rio Coyolapan, 
and Rio Coyolate. The fruiting panicles are very large, heavy, and 
pendent, producing large numbers of the small fruits, of characteristic 
appearance. These are boiled with sugar and eaten. The amount 
of flesh is scant, and the fruits are a very poor food, but great quan- 
tities of them thus prepared are offered for sale in most of the markets 
and must find ready customers. The kernels of the seeds, too, are 
sometimes eaten, at least by children. Boys at Huehuetenango 
cracked them by placing the nut between cobblestones, then throw- 
ing rocks at it. They said they didn't crack the nuts with a stone 
held in the hand because they were too hard to be broken that way. 
The hard part of the nut is used for making finger rings, rosary beads, 
and other articles. The fresh inflorescences often are used as deco- 
rations, and bring a good price for this purpose in regions where the 




FIG. 35. Acrocomia mexicana. a, Habit, b, Portions of leaf (X Y%). c 
Fruit, from below (X V\}. d, Fruiting cluster (X Y*). e, Spathe (X V*). 



203 



204 FIELD IANA: BOTANY, VOLUME 24 

palm is not native. The fruits are much eaten by cattle and other 
stock. In cavities in felled trunks there collects a quantity of sap 
which is drunk when beginning to ferment, or later, when it some- 
times is intoxicating. It is a favorite beverage among the vaqueros 
of the Pacific coast. This vino de palma is well known in most parts 
of Central America where Acrocomias are native. 



ASTEROGYNE Wendland 

Reference: M. Burret, Bot. Jahrb. 63: 140. 1930. 

Small unarmed plants, the caudex slender, densely annulate; leaves simple, 
oblong, bifid, usually entire, the segments acuminate, plicate-nerved; spadix long- 
pedunculate, once-branched, the branches subfasciculate, strict, rigid, stout; 
spathes 2-4, subpersistent; bracts small, continuous with the orifice of the pit of 
the rachis, the bractlets scale-like; flowers small, glumaceous, unisexual, sunken 
in spirally arranged pits in the rachis, in groups of 3-1, the central flower pistillate 
(or abortive) maturing after the lateral staminate flowers; staminate flower asym- 
metric, the sepals obovate-oblong, carinate, imbricate, the petals concave, valvate; 
stamens 6, the filaments connate, free at the apex, the anthers pendulous; pistillate 
flowers compressed, the perianth unchanged after anthesis, the sepals oblong, cari- 
nate, imbricate, the corolla trifid, the lobes stellately recurved, imbricate, the tube 
adnatc to the disk and with it circumscissle above the base; ovary 3-celled, the style 
apical, elongate, the stigmas exserted; fruit ellipsoid-oblong, subcompressed, nar- 
rowed at each end, 1-seeded, the scar of the style basal, the pericarp thin, juicy, 
fibrous within, the fibers slender, adherent to the endocarp; seed oblong, erect, the 
testa smooth, brown, the hilum basal, the endosperm uniform, the embryo basal. 

One other species is known, in Colombia. 

Asterogyne Martiana Wendland ex Hook. f. in Benth. & Hook. 
Gen. PL 3: 914. 1883. Geonoma Martiana Wendl. Linnaea 28: 342. 
1856. G. trifurcata Oerst. Vid. Medd. Kjoebenhavn 1858: 34. 1859. 
Capoca (Izabal). Figure 36. 

Dense wet mixed lowland forest, at or little above sea level; 
Izabal. British Honduras, along the Atlantic coast to Panama; 
Colombia. 

Plants small, often forming clumps or thickets, commonly acaulescent or 
nearly so but sometimes with a caudex 1.5 meters high and 4-5 cm. thick; leaves 
numerous, dull deep green, 1.3 meters long or shorter, deeply bifid at the apex, 
cuneately narrowed at the base, with about 40 nerves on each side; spadices 60- 
70 cm. long, the peduncle 40-50 cm. long, the rachis short, 4 cm. long or less, the 



FIG. 36. Asterogyne Martiana. a, Leaf (X X A)- b, Fruit (X 4). c, Fruiting 
branch (X M)- d, Staminate flower (X 11). 



206 FIELDIANA: BOTANY, VOLUME 24 

branches 3-6, dull pale red, 10-20 cm. long; pits of the rachis 7-ranked, 9-10 mm. 
apart, the flowers white, fragrant; fruits spreading from the rachis, bluish black 
when ripe, smooth; seeds oval-ellipsoid, 6-9 mm. long, 4-5 mm. broad. 

Called "monkey- tail" (British Honduras), "pacuquilla" (Hon- 
duras), "pato de gallo," "suita" (Nicaragua). In general appearance 
the plant suggests a Geonoma, but from the Guatemalan species of 
that genus it is distinguished at once by its simple leaves, as well as 
by the fasciculate spadix branches, the rachis of the spadix being 
very short or almost obsolete. 

ASTROCARYUM Mey. 

Rather small palms, the slender trunk armed with irregularly spaced, long, 
stout, compressed spines; leaves large, irregularly pinnatisect, the petiole short, the 
sheath short and open; spathes 2, borne at the base of the leaf cluster, white-lanate, 
covered with slender needle-like spines; spadix rather small, branched, the flowers 
monoecious in the same spadix, the staminate portions of the branches soon wither- 
ing and falling, the basal flower of each branch pistillate, larger than the staminate 
flowers, the upper flowers all staminate and densely crowded; pistillate calyx 
cupuliform, the corolla urceolate, tridentate, bearing within 6 small, free or united 
staminodia; ovary ovoid, 3-celled, the 3 stigmas short, the cells 1-ovulate; stami- 
nate calyx small, triangular, the 3 petals short-stipitate, thick and fleshy, valvate; 
stamens 6, the filaments thick, the anthers dorsifixed, linear, sagittate at the base; 
fruit rather large, rostrate, 1-seeded, ovoid, spiny, the pericarp fleshy, fibrous 
within, the endocarp thin-osseous, 3-pored at the apex, the pores stellate-fibrous, 
the fertile pore apical; embryo apical; endosperm simple. 

The genus consists of about 48 species. 

Astrocaryum mexicanum Liebm. Dansk. Vid. Selsk. Forh. 
1845: 8. 1846 (type from Oaxaca). A. Warscewiczii Koch & Fint. 
Wochenschr. 1: 297. 1858. Bactris Cohune Wats. Proc. Amer. Acad. 
21: 467. 1886 (type from "Chocon forests," Izabal). A. Cohune 
Standl. Trop. Woods 21: 25. 1930. Hexopetion mexicanum (Liebm.) 
Burret, Notizbl. Bot. Gart. Berlin 12: 156. 1934. Lancetilla; Guis- 
coyol; Warree cohune, Cohune (English-speaking negroes of North 
Coast and British Honduras). Figure 37. 

Common and often abundant in dense wet mixed lowland forest 
of the northern departments, usually on well-drained land, chiefly on 
hillsides, 400 meters or lower; Pete"n; Alta Verapaz; Izabal. South- 
ern Mexico (Oaxaca, Tabasco); British Honduras; Atlantic lowlands 
of Honduras. 

Trunk slender, 1.5-5.5 meters high, 2.5-4.5 cm. in diameter, armed with very 
numerous, spreading, stout spines, these compressed and 2-edged, blackish, 3-5 cm. 
long or longer, arranged in more or less regular whorls; leaves a meter long or larger, 




FIG. 37. Astrocaryum mexicanum. a, Leaf (X VT). b, Inflorescence with 
spathe (X V*)- c, Longitudinal section through fruit (X %). d, Fruit (X %). 
e, Portion of spiny trunk (X V*). /, Fruiting cluster (X M)- 



207 



208 FIELDIANA: BOTANY, VOLUME 24 

glaucous beneath, the petiole and rachis more or less tomentose and armed with 
long needle-like spines, the segments broad or narrow, very unequal, as much as a 
meter long, sparsely aculeolate on the margins; spathes 20-30 cm. long, caudate- 
acuminate; staminate calyx densely lanate, the petals lanceolate, united at the 
base, 4 mm. long; pistillate calyx and corolla aculeolate, coriaceous, the corolla 
12-16 mm. long, the calyx almost half as long; fruits very numerous in each spadix, 
obovoid, about 5 cm. long, 2-2.5 cm. broad, rostrate, densely covered with short 
slender spines; seed obovoid, acute at the base, rounded at the apex, 2.5 cm. long. 

This unpleasantly armed palm often forms a dense undergrowth, 
especially on hillsides, in the dark forests of the Atlantic lowlands, 
the very numerous, sharp-tipped spines being a constant menace to 
any one pushing through the thickets. Watson states that the name 
"warree cohune" is given because of the resemblance of the spiny 
covering of the fruit to the bristles of the common peccary (called 
"warree" by the negroes of the Atlantic coast). He states also that 
the kernels of the fruits are edible, with a flavor similar to that of 
coconut. The common Spanish name "lancetilla" alludes to the 
lance-like spines. 

BAGTRIS Jacquin 

Reference: Burret, Repert. Sp. Nov. 34: 167, 237, 241. 1934 (in- 
cluding Guilielma Martius and Pyrenoglyphis Karsten). 

Low or tall palms, abundantly armed with short or long spines, the stems soli- 
tary or forming dense clumps or colonies, the stems or trunk annulate; leaves 
terminal or scattered along the upper part of the stem, equally or unequally pin- 
natisect, the sheath elongate, spiny; spathes 2, longer than the spadix, cymbiform 
or fusiform; spadix simple or simply branched, inserted among the leaves; flowers 
monoecious in the same spadix, sessile, the lower ones ternate with the middle 
flower pistillate, or the sexes irregularly scattered; staminate calyx annular, urceo- 
late, or 3-parted; stamens 6, 9, or 12, included, the filaments subulate, the anthers 
linear, affixed by the bifid base, erect; pistillate calyx various, the corolla longer 
than the calyx or of the same length, tridenticulate at the apex, the staminodia 
free and dentiform, or united in a ring; ovary 3-celled, the 3 stigmas short, sessile, 
finally recurved; fruit ovoid, subglobose, or oblong, 1-celled and 1-seeded, the stig- 
mas terminal, the pericarp hard and almost ligneous or fleshy and juicy, the endo- 
carp osseous, 3-pored near the apex or above the middle; seed pendulous below the 
apex of the cell, the raphe reticulate, the endosperm uniform, corneous, the embryo 
opposite one of the pores. 

Nearly 200 species are known, distributed almost throughout 
tropical America but mostly in South America. A few species are 
found in Mexico, and more occur farther south in Central America. 
It is probable that others may be found in Guatemala, but they have 
not been collected. 

Many of the species are more offensively armed, perhaps, than 
any other Central American plants, and that is saying a great deal. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 209 

Areas thus protected furnish admirable hideouts for all sorts of ani- 
mals, particularly peccaries, agoutis, and other animals that feed on 
the fruits. The fruits, indeed, often are eaten by people, but they 
furnish a sorry sort of food. Those of some species do have a small 
amount of acidulous pulp that is not unpalatable, but others, which 
are eaten in extreme cases, have little that can be consumed except 
the very hard kernels, and these are eaten usually in the green and 
immature state, if at all. An exception is the pejibaye, B. Gasipaes, 
which is cultivated for its edible fruit. 

Fruit purple at maturity, 3.5-4 cm. long; pistillate flowers with staminodes united 
in a ring and forming a conspicuous crenate cupule within the corolla in fruit. 

B, balanoidea 
Fruit red or yellow at maturity; pistillate flowers with inconspicuous free dentiform 

staminodes. 
Fruit large, fleshy, edible; endocarp with numerous flat fibers radiating from the 

pores B. Gasipaes 

Fruit small, about 1.5 cm. high and broad; endocarp lacking radiating fibers. 

B. trichophylla 

Bactris balanoidea (Oerst.) Wendl. in Kerch. Palm. 233. 1878. 
Augustinea balanoidea Oerst. Vid. Medd. Kjoebenhavn 1858: 39. 
1859. Pyrenoglyphis balanoidea Karsten, Fl. Colomb. 2: 142. 1869. 
Guiscoyol; Viscoyol; Huiscoyol; Pahuac (Pete"n). Figure 38. 

Dry to wet thickets or forest, often in swampy ground, mostly at 
250 meters or lower; Pete"n; Izabal; Santa Rosa; Escuintla; Retalhu- 
leu; probably in all the Pacific coast departments. Tabasco and 
probably elsewhere in southern Mexico; doubtless in British Hon- 
duras, and extending southward to El Salvador and Panama. 

Plants slender, the stems cane-like, cespitose, and usually forming dense, small 
or large clumps or thickets, generally 5 meters high or lower, the stems 2.5-5 cm. 
in diameter, densely armed with long and short, blackish spines, or the older stems 
almost naked; leaves evenly pinnatisect, mostly 1-1.5 meters long, dark green 
above, dull pale green beneath, wide-spreading and more or less decurved, the 
slender petioles 45-60 cm. long, ferruginous-pubescent and armed with spines 2.5- 
7.5 cm. long; leaf segments 30-40 pairs, subopposite or the lower ones alternate, 
2-5 cm. apart, 25-45 cm. long, puberulent beneath, the margins setulose; spathe 
20-35 cm. long, densely spiny; spadix with 10-12 simple branches; fruit cream- 
colored, probably becoming purple, 3.5-4 cm. long and 1-2.5 cm. thick, obtuse or 
rounded and apiculate at the apex; calyx persistent, 8-11 mm. high; endosperm 
surrounded by coarse hair-like fiber, about 3 cm. long, black. 

Burret considers it doubtful that this differs from B. major (Jacq.) 
Karst., which ranges from Panama to Venezuela. The small spadices 
of this palm are sometimes used in Guatemala for decorating nacimi- 
entos, according to Professor Ulises Rojas. 




a 



FIG. 38. Bactris balanoidea. a, Apical, median, and lower portions of leaf 
( X V*)- b, Fruit, side view ( X %). c, Fruit, ventral view ( X M). d, Entire leaf 
(X Vio)- e, Fruiting branch with spathe (X l /i). 



210 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 211 

Bactris Gasipaes HBK. Nov. Gen. & Sp. 1: 302, pi. 700. 1816. 
GuilielmaGasipaes L. H. Bailey, Gentes Herb. 2: 187. 1930. G. utilis 
Oerst. Vid. Medd. Kjoebenhavn 1858: 46. 1859. Pejibaye. 

Native from Costa Rica to Colombia, and perhaps southward to 
Peru; planted occasionally in Guatemala at low and middle eleva- 
tions, particularly in the North Coast, also in Escuintla and at 
Antigua. 

Trunk sometimes 8 meters high or more; fruits either red or yellow at maturity. 

Apparently only a few plants of the pejibaye are in cultivation in 
Guatemala but in Costa Rica in certain regions they are abundant, 
both wild and cultivated. It is, however, unknown whether the 
apparently wild plants really are so, or only remnants of former 
plantations. In Costa Rica the palm is sometimes called "chonta," 
a word of South American (Quechua?) origin. In that country the 
pejibaye is one of the most highly valued plants because of its abun- 
dant fruits, which are much sought in the markets during their brief 
season. When boiled, the flesh of the fruit is mealy and sweet, in 
flavor and consistency somewhat suggesting boiled chestnuts (Cas- 
tanea) or sweet potato. The very hard wood was used formerly by 
the Indians of Costa Rica for fashioning bows, arrow points, and 
other articles. Like other palms, this is an ornamental one for culti- 
vation, the chief objection to it being the too great abundance of 
offensive spines on the trunk, which makes it unsuitable for planting 
too close to dwellings. 

Bactris trichophylla Burret, Repert. Sp. Nov. 32: 113. 1933; 
34: 214. 1934. Guiscoyol; Huscoyol. Figure 39. 

Moist or wet thickets, often in wooded swamps, 900 meters or 
lower; Pete'n; Alta Verapaz; Izabal. British Honduras, the type 
from Stann Creek Valley, at 300 meters, W. A. Schipp S368. 

Plants rather large and as much as 8 meters high, cespitose and often forming 
dense clumps or thickets, the stems 3-6 cm. in diameter; petioles 60 cm. long or 
more, furfuraceous-tomentose or glabrate, armed below with slender but stiff, 
blackish spines 4-7 cm. long; leaf blades 1.5-2 meters long, the rachis densely 
spiny below; leaf segments about 25 on each side, grouped, linear, mostly 30-60 cm. 
long and 2-4 cm. wide, acuminate, 2-3-nerved, deep green above, somewhat paler 
beneath and pubescent; upper spathe 24 cm. long, narrow-acuminate, fuscous- 
leprose, not at all tomentose, armed with slender spines 5-10 mm. long; peduncles 
about 11 cm. long, pale-tomentose, densely spiny, the branched portion 16 cm. 
long, the branches about 40 and 8-12 cm. long, fuscous-furfuraceous; pistillate 
calyx 3.5 mm. broad, with 3 very short, acute teeth, the corolla obscurely 3-den- 
tate; ripe fruit depressed-globose, mammillate, 1.5 cm. high and broad, glabrous; 
fruiting perianth 1 cm. broad. 




S.H. Grove 



212 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 213 

This plant and Bactris balanoidea are much alike in the field and 
have in common their most outstanding and offensive characters, 
which commonly attract more attention than the differences by 
which botanists distinguish them. Both plants are known among 
the native people, who naturally do not distinguish the plants, by 
the name "huiscoyol" or "giiiscoyol," a Nahuatl term signifying 
"spiny palm." They are abundant in the lowlands of Guatemala. 

BRAHEA Martius 

References: H. H. Bartlett, The genus Brahea, Carnegie Inst. 
Wash. Publ. 461: 29-31. 1935; L. H. Bailey, Notes on Brahea, Gentes 
Herb. 4: 119-125. 1937; L. H. Bailey, Brahea, and an Erythea, Gentes 
Herb. 6: 177-197, /. 87-100. 1943; H. E. Moore, Some American 
Corypheae, Gentes Herb. 8: 219-222. 1951. 

Erect or decumbent, sometimes acaulescent, solitary or cespitose, medium- 
sized hermaphrodite palms, the trunk fibrous-roughened below, clothed with a 
shag of deciduous leaf sheaths near the apex; leaves palmate, the sheath soft 
fibrous, unarmed; petiole with entire or spinose-dentate margins, ligule present at 
base of the blade above, lacking below; blades flabellate to nearly orbicular in 
outline, very shortly costapalmate, regularly divided nearly to the middle or be- 
yond into 1-nerved segments bifid at the apex; spadices interfoliar, equaling or 
exceeding the leaves, erect or arcuate, slender with several once- to thrice-branched 
primary divisions, the spathes tubular, sheathing the peduncle, primary and sec- 
ondary branches; flowers small, solitary, sessile on tomentellous branchlets or im- 
mersed in thick tomentose branchlets; sepals 3, free, imbricate; petals 3, free or 
shortly united with stamen filaments at the base, valvate, foveolate; stamens 6, 
the filaments united in a ring at the base and adherent to the corolla, attenuate 
above; carpels 3, glabrous above, tomentose below and prominently so as fruit 
develops, lightly connate below, the styles connate in a conic beak stigmatic at the 
apex, the ovule basal and erect; fruit ellipsoid, the stigmatic scar subapical, abor- 
tive carpels basal, exocarp smooth or tomentose when young, mesocarp fleshy, 
endocarp crustaceous, fragile; seed ellipsoid or globose, the raphe narrow, endo- 
sperm homogeneous with an intrusion of the integument below the raphe, embryo 
lateral. 

A small genus of perhaps eight species, all except the following 
confined to Mexico. 

Flowers sessile or slightly elevated on slender tomentellous branchlets; leaves con- 
spicuously waxy-glaucous below, the petioles unarmed B. prominens 

Flowers immersed in thickish densely tomentose branchlets; leaves not waxy- 
glaucous below, the petioles spinose-dentate along the margins B. dulcis 



FIG. 39. Bactris trichophylla. a, Leaf (X l /\o). b, Portion of trunk with 
fruiting branch and spathe (X */) c, Fruit (X 



214 FIELDIANA: BOTANY, VOLUME 24 

Brahea dulcis (HBK.) Martius, Hist. Nat. Palm. 3: 244. 1838. 
Corypha dulcis HBK. Nov. Gen. et Sp. 1: 300. 1816. Apak (Hue- 
huetenango). 

Open rocky hillsides, 800-1200 meters; Huehuetenango; Quezal- 
tenango. 

Plants to 6 meters high or somewhat more, the stems solitary or sometimes 
cespitose; leaves 50-70 cm. across, divided into 30 or more segments 2-3 cm. wide, 
the petioles to 75 cm. long or more, margined with small teeth 2-4 mm. high; 
blades green; inflorescences to 1 meter long or more, with several twice- to thrice- 
branched primary divisions, the ultimate branchlets densely yellow-brown tomen- 
tose and thickish, 5-15 cm. long, with solitary flowers imbedded in the tomentum; 
flowers about 2 mm. high, the sepals scarcely prominent, petals prominently canes- 
cent at the base and glabrous above; fruit to 17 mm. long when fresh but smaller 
when dry, ellipsoid, yellowish-brown. 

Brahea prominens L. H. Bailey, Gentes Herb. 6: 192. 1943. 
Palma. 

At 900-1600 meters; Alta Verapaz (limestone cliffs above Trece 
Aguas, Cook & Doyle 3}; Huehuetenango (?). 

Trunk erect, 10 meters high or less, as much as 30 cm. in diameter at the base; 
leaves about a meter broad, orbicular, silvery beneath, light green above, glabrous, 
divided to the middle or lower into about 50 narrow segments, the margins with 
few, very short filaments; midrib of the rachis continued about 15 cm. on the lower 
surface of the blade; petioles very long (about 180 cm.), glaucous, not toothed; 
spadix very long and in fruit exceeding the leaves and hanging beneath them, the 
almost cylindric, vaginate peduncle longer than the fruiting portion of the inflores- 
cence, much branched, the ultimate branchlets 5-8 cm. long, slender, tomentellous; 
flowers 2-3 mm. long, sessile or slightly elevated on the branchlets; fruit straight 
or flattened on the ventral side, 16-17 mm. long, 10 mm. thick when fresh, smaller 
when dried, light dull yellow or greenish yellow, the outer pulp scant, the apex 
with a short oblique point, the cupule very small, with acute corolla lobes. 

The species of Brahea are still imperfectly known. Specimens 
from Guatemala agree with B. prominens but it remains to be seen 
whether the species is distinct from the earlier B. calcarea Liebmann. 



CALYPTROGYNE Wendland 
Reference: M. Dammer, Bot. Jahrb. 63: 131-140. 1930. 

Plants unarmed, usually small and low, sometimes stoloniferous, the caudex 
short or elongate, annulate below; leaves terminal, unequally pinnatisect, the seg- 
ments few, narrow or broad, long-acuminate, often falcate, plicate, with numerous 
nerves, the petiole short, the sheath short, open; spadices simple or branched from 
the base, long-pedunculate; spathes 2, narrow, the lower one much shorter than 
the peduncle, cleft at the apex, the upper deciduous, elongate, cleft its whole length; 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 215 

flowers monoecious, inserted in pits in the rachis of the spadix, the lower bract 
forming a broad lip projecting from the pit, the flowers ternate, the middle one 
pistillate, rarely solitary or binate, the perianth glumaceous; staminate flower sub- 
symmetric, compressed, the 3 sepals oblong or lanceolate, concave, 2 or all of them 
dorsally carinate or winged; petals connate at the base into a stipe, ovate or spatu- 
late, valvate; stamens 6, the filaments connate to form a tube, their apices free, 
the anthers sagittate, basifixed; pistillate flower broadly ovoid and strongly com- 
pressed, the perianth unchanged after anthesis, the petals valvate; ovary 3-celled, 
the style central, short or elongate, the stigmas recurved; fruit oblong or obovoid, 
1-seeded, the scar of the style basal, the pericarp fleshy and fibrous; seed oblong or 
subglobose, slightly compressed, the hilum basal, the branches of the raphe incon- 
spicuous, the endosperm uniform (not ruminate), the embryo basal. 

About 11 species are known, ranging from southern Mexico to 
Brazil and Peru. A few others are found in other parts of Central 
America. 

Calyptrogyne Donnell-Smithii (Dammer) Burret, Bot. Jahrb. 
63: 133. 1930. Geonoma Donnell-Smithii Dammer, Bot. Jahrb. 36, 
Beibl. 80: 32. 1905. Capoca; Guiscoyol (doubtless an erroneous name, 
belonging properly to Bactris and Pyrenoglyphis) ; Coroz. Figure 40. 

Dense wet mixed lowland forest, sometimes in Manicaria swamps, 
600 meters or less, mostly at or near sea level; Izabal; perhaps en- 
demic but to be expected in British Honduras and Honduras. 

Plants small, acaulescent or with a very short caudex; leaves 1 meter long or 
more, the petiole short, 40 cm. long or usually much shorter, the blades dull deep 
green, unequally pinnatisect, the segments linear or broader, 35-45 cm. long, the 
blade with about 24 nerves on each side; inflorescence simple, the peduncle very 
slender, greatly elongate, stiff but flexible; lower spathe 26 cm. long or more; rachis 
of the spadix dull salmon-red to dark red, about 25 cm. long and 7 mm. thick, gla- 
brous, the pits 7-ranked; staminate sepals 4.5 mm. long, the pistillate 4 mm. long; 
fruits ascending along the rachis (not divaricate), globose-oval, black at maturity, 
about 8 mm. long when dried, larger when fresh. 

This palm has been reported from Guatemala under the name 
Geonoma Olfersiana Klotzsch, which belongs to a true Geonoma native 
in Brazil. 

CARYOTA L. 

Plants unarmed, monocarpic, the trunk stout and tall, annulate; leaves very 
large, disposed in a terminal, often somewhat elongate crown, bipinnatisect, the 
numerous segments half-fan-shaped or wedge-shaped, dentate at the apex; margins 
of the leaf sheaths fibrous; spadices often alternately staminate and pistillate, short- 
pedunculate, the branches elongate and pendulous; spathes 3-5, incomplete, tubu- 
lar; flowers rather large, monoecious in the same or different spadices which are 
inserted among the leaves, ternate with the middle flower pistillate, or the stami- 
nate flowers solitary, the spadices fastigiately branched; staminate sepals rounded, 




FIG. 40. Calyptrogyne Donnell-Smithii. a, Inflorescence (X %). b, Leaf 
(X Yz). c, Unopened flower (X 5). d, Staminal portion of flower (X 15). e, 
Fruit with attachment to rachis (X 27). 

216 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 217 

coriaceous, imbricate, the petals linear-oblong, valvate; stamens numerous, the 
anthers linear, affixed by the bifid base; pistillate flowers smaller than the stami- 
nate, subglobose, the perianth scarcely accrescent after anthesis, the sepals rounded, 
broadly imbricate, the petals coriaceous, rounded, connate at the base, valvate 
above; ovary 3-celled, the stigma trilobate, sessile, the ovules basal, erect; fruit 
globose, 1-2-seeded, the stigma terminal, the pericarp thin, coriaceous; seed sub- 
globose, the hilum basal, the branches of the raphe sunk in channels, the endo- 
sperm ruminate, the embryo dorsal, excentric. 

The genus consists of about a dozen species in tropical Asia and 
East Indies. 

Caryota mitis Loureiro, Fl. Cochinch. 569. 1790. Cola de 
pescado. 

Native of southeastern Asia; occasionally planted for ornament 
in Guatemala, at low or middle elevations. 

Trunk often 5-8 meters high or more and 10-13 cm. in diameter; petioles, leaf 
sheaths, and scapes scurf y-villous; leaves 1-3 meters long, the numerous pinnae 
very obliquely cuneiform, irregularly dentate, the upper margin acute. 

Caryota urens L. Sp. PL 1189. 1753. 

Native of southeastern Asia and Malaysia; like the preceding 
species, planted for ornament. 

Trunk thicker and taller than in C. mitis, becoming 25 meters tall and 45 cm. 
in diameter; leaves 5-6 meters long and 3-3.5 meters wide, the pinnae 1.5-2 meters 
long, recurved and drooping, the leaflets very obliquely truncate, 10-20 cm. long, 
acutely serrate, the upper margin caudately produced. 

In Asia called "wine palm" and "toddy palm," being the source 
of a palm wine that is highly esteemed there. Both species are 
known in Florida, where they are sometimes planted, by the name 
"fishtail palm." They are easy of recognition because of their bipin- 
natisect leaves, which are large and broad, numerous, and handsome 
in appearance. In Guatemala these palms are seen mostly in the 
lowlands, where they often are planted in the parks. Because of the 
appearance of the leaf segments, they are often referred to as "palma 
comida por las ratas." 

CHAMAEDOREA Willdenow 
Reference: Burret, Notizbl. Bot. Gart. Berlin 11: 724. 1933. 

Plants unarmed, erect or procumbent, rarely scandent, the stems solitary or 
cespitose, very slender, green, annulate; leaves simple and bifid or rarely rounded 
at the apex, or variously pinnatisect, the segments broad or narrow, few or numer- 
ous, often somewhat oblique or sigmoid, acuminate, plicate-nerved, the petiole 



218 FIELDIANA: BOTANY, VOLUME 24 

usually slender and cylindric, the sheath short or elongate, tubular; spadices borne 
among or below the leaves, simple or paniculately branched, the peduncle short or 
elongate, the branches sparsely or densely flowered; spathes 3 or more, elongate 
and sheathing the peduncle, cleft at the apex, coriaceous or membranaceous, more 
or less persistent; flowers dioecious, sessile or subimmersed in the spadix, small or 
minute, ebracteate and ebracteolate; staminate calyx cupuliform, entire, 3-lobate, 
or 3-parted, the 3 petals free or variously connate, the lobes or segments valvate; 
stamens 6, the filaments short, broad or subulate, the anthers dorsifixed, included, 
didymous or oblong; pistillate calyx like that of the staminate flower, the petals 
usually connate, valvate or imbricate; ovary 3-celled, the stigmas minute, recurved, 
the ovules solitary, basal, erect; fruit small, of 1 or sometimes 2-3 carpels, globose 
or oblong, the stigmas basal, the pericarp coriaceous or carnose, scarcely fibrous; 
seed erect, globose or ellipsoid, the small hilum basal, the branches of the raphe 
obscure, the endosperm cartilaginous, uniform, the embryo basal or dorsal. 

About 100 species are known, ranging from central Mexico to 
Brazil and Bolivia. A good many others are found in southern Cen- 
tral America, especially in Costa Rica (16 species) and Panama (19 
species) . Although Burret has done much to establish some order in 
the genus, most of the species still are known from only a few speci- 
mens, and their classification is still far from satisfactory. Burret 
recognizes as distinct from Chamaedorea the genera Collinia (C. ele- 
gans), Eleutheropetalum (Chamaedorea Ernesti-Augustii and its rela- 
tives), and Dasystachys (Chamaedorea Deckeriana). The differences 
stressed, however, do not seem of generic importance in a genus 
where variation in floral morphology is considerable. It therefore 
seems more practical, at least, and probably more in accordance with 
general taxonomic custom, to treat those genera as synonymous 
with Chamaedorea. 0. F. Cook has gone still farther, separating 
from Chamaedorea as genera numerous forms that would be treated 
by most botanists as species, an extremity of splitting that has not 
been surpassed even in the Cactaceae or Leguminosae. 

The Chamaedoreas are of considerable economic importance 
throughout Central America, but in Guatemala more than else- 
where. In the markets there are sold vast quantities of the un- 
opened staminate inflorescences (the pistillate ones are not used), 
that is, the twisted and folded spadix enclosed by the closely ap- 
pressed, green spathes, the whole looking like a green ear of maize 
enclosed in its husks. The spadices are white or whitish, tender, 
and when served on the table are of attractive and agreeable flavor. 
They constitute, indeed, an excellent vegetable and one that is very 
popular locally. The spadices of some species are too bitter to be 
eaten, and all or most of them have a slight bitterness that is rather 
agreeable. Sometimes they are served raw as a salad but more often 
they are cooked with eggs, or prepared in other ways, much as aspara- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 219 

gus is treated. In some regions, particularly in the Occidente, there 
is prepared from Chamaedorea a quite different dish or vegetable, 
called "bojon." This consists of the pith of the younger stems, and 
perhaps includes at times the leaf buds. It is eaten either raw, some- 
times as a salad, or cooked in various fashions. The senior author 
has had it served on the table baked, and it proved to be succulent 
and well flavored, much better, in fact, than even the ordinary pacaya. 

The spadices of different species of Chamaedorea as they appear in 
the markets differ greatly in appearance. The pacayas (the name 
"pacaya" is applied to both the plant and its edible spadices, or the 
plant is sometimes called "pacayo") of Cobdn (C. Tepejilote) are 
much larger and finer than those of other localities, and when piled 
on the ground resemble green ears of field corn. Such spadices, 
naturally, give a larger amount of edible matter than those of smaller 
size, and the Alta Verapaz pacayas on that account are shipped to 
other parts of the country. We have seen large trucks loaded with 
them starting for the Guatemala market. In Alta Verapaz the pa- 
caya has been a cultivated plant probably for centuries, and the 
plants have increased in size as the result of cultivation, being much 
taller and stouter than plants of the same species found in the moun- 
tains. Some of them are taller than the one-storied houses. While 
the large pacaya is referred to commonly as the pacaya of Coban, 
the plants actually are grown mostly far from that city, and they are 
especially abundant in Tactic and San Cristobal, where there are 
veritable forests of them in some of the small fincas. 

Some species of Chamaedorea have spadices so small that they are 
worthless for food, but most of those eaten in Central America are 
much smaller than those of C. Tepejilote. Almost all the species may 
be found in cultivation at times in patios or gardens, and all of them 
are ornamental. The larger plants are valued more for their food 
value than for ornamental purposes, and many patios have clumps 
of the plants from which the inflorescences are gathered for the table. 
The spadices are produced almost if not quite throughout the year, 
and are one of the vegetables that may be procured at all seasons. 
The larger leaves are much used for decorations on festive occasions, 
both inside and out of doors, and they are especially in favor for form- 
ing arcos across streets and roads. The juice of ripe Chamaedorea 
fruits is intensely irritating to the skin, causing burning and itching 
sensations. Mischievous boys sometimes push through crowds of 
people, rubbing the juice on their arms, much to the discomfort of 
persons so molested. 



220 FIELDIANA: BOTANY, VOLUME 24 

The general name in Guatemala for species of Chamaedorea is 
pacaya, and this term appears in many geographic names, especially 
that of the Volcan de Pacaya. The name "tepejilote," current in 
Mexico, is seldom heard in Guatemala, but it is said to be applied to 
one of the small plants with very bitter spadices. Other names are 
"capuca," "chichicuilote," "bojon," and "molenillo." 

Flowers of both sexes with petals united, opening through a terminal 3-angled pore, 
the corolla in pistillate flowers splitting into 3 rather strongly nerved equal 
lobes in fruit; pistillodes of staminate flowers expanded into a broad 6-angled 
cap with anthers adherent to the column below C. elegans 

Flowers of both sexes with petals distinct above a sometimes shortly united base, 
or with staminate petals connate at the apex and the corolla opening by 
lateral slits. 

Petals bright orange at maturity in both sexes, the pistillate valvate at anthesis 
(probably lightly imbricate in bud basally) with a thin basal half and a 
fleshy cucullate upper half, the latter deciduous in fruit; pistillode of stami- 
nate flowers expanded into a broad 6-lobed cap, the anthers deeply bifid at 
the apex; leaves simple; inflorescence long-pedunculate. 

C. Ernesti-Augustii 

Petals yellow, green, or brownish at maturity in both sexes, the pistillate imbri- 
cate and persistent in entirety about the fruit (except probably in C. brachy- 
poda); pistillode truncate or attenuate, scarcely much expanded at the apex; 
leaves and inflorescences various. 

Staminate flowers contiguous, densely crowded in 4-7 rows along the branches 
of the inflorescence and lightly immersed in shallow elliptic depressions; 
pistillate flowers with the calyx either very small and annular or cupular 
and nearly as long as the petals. 

Pistillate inflorescence with few to many slender branches, the flowers with 
a very low annular calyx and not densely crowded; staminate flowers 
with the calyx very low and annular, not prominent in bud. 

C. Tepejilote 

Pistillate inflorescence spicate, furcate, or with few thick branches, the 
flowers very densely crowded, with cupular calyx nearly half as long 
as the petals; staminate flowers with the calyx well developed, cupular 
and prominent in bud. 

Pistillate inflorescence spicate or furcate C. Arenbergiana 

Pistillate inflorescence with about 8 branches C. Casperiana 

Staminate flowers distinct and separated, at least in bud (at anthesis dense in 
C. adscendens) ; pistillate flowers not densely crowded and with a well- 
developed imbricate or 3-lobed calyx. 
Petals of staminate flowers connate at the tips, the corolla opening by 

lateral slits; staminodes lacking in pistillate flowers. 
Perianth of both sexes nerveless or essentially so; leaves simple; inflores- 
cences with 1-6 branches; fruit maturing black . . .C. geonomaeformis 
Perianth of both sexes very strongly nerved when dry; fruit orange or 
yellow, at least when immature, in some species maturing purple- 
black. 

Flowers of both sexes essentially superficial in slight depressions along 
the branches of the inflorescence, the sepals generally united and 
scarcely imbricate-lobed in bud; inflorescence of both sexes with 5 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 221 

(rarely 4) or more branches; petiole elongate, usually exceeding the 
leaf sheath in length. 
Plants of Alta Verapaz at elevations of 1350-1600 meters; fruit 

unknown; see discussion of species C. micrantha 

Plants of western Guatemala, or of low elevations (mostly less than 

300 meters) in eastern Guatemala. 
Fruit sickle-shaped, acute; branches of the staminate inflorescence 

few (4-7) '. C. falcifera 

Fruit globose or subreniform and obtuse; branches of the staminate 

inflorescence numerous. 
Fruit globose; leaves small with generally 5 pinnae on each side 

of a rachis 30-40 cm. long C. Aguilariana 

Fruit subreniform, obtuse; leaves large with usually 7-8 pinnae 

on each side of a rachis 45-65 cm. long C. neurochlamys 

Flowers of both sexes distinctly sunken in prominent depressions along 
the branches of the inflorescence; inflorescences of both sexes spi- 
cate, furcate, or trisect; staminate sepals nearly completely imbri- 
cate in bud; petioles short, seldom equaling, very rarely exceeding 
the sheath in length. 

Leaves thin, simple or generally pinnate below broad terminal lobes, 
with 1-3 secondary nerves clearly evident between the primary 
nerves of the terminal lobes, the rachis generally 8.5 cm. long or 
more, very rarely only 6 cm. long; pistillate flowers prominently 
sunken, the pits deep in fruit; caudices erect, solitary, with a 

cluster of slender prop or prop-like roots C. Rojasiana 

Leaves rather thick, simple, the secondary nerves not evident, the 
rachis generally 5 cm. long or less; pistillate flowers lightly 
sunken, the depressions shallow in fruit; stems decumbent and 
rooting at the nodes, then erect, plants possibly rhizomatous 

and colonial C. simplex 

Petals of staminate flowers free at the apex, the perianth of both sexes 
nerveless or with few nerves when dry; fruit green when immature, 
maturing black or glaucous. 

Branches of the inflorescence undulate, especially in bud, the flowers of 
both sexes laterally compressed, longer than broad, slightly to dis- 
tinctly impressed in curves of the branches. 

Pistillate flowers with staminodes, the calyx 3-lobed C. nubium 

Pistillate flowers lacking staminodes, the calyx of essentially imbricate 

sepals C. Skutchii 

Branches of the inflorescence not undulate, though sometimes ridged and 

angled; flowers essentially superficial, not laterally compressed. 
Inflorescences of both sexes rather profusely branched, generally with 
10 or many more branches (only 3-6 in pistillate plants of C. vul- 
gata but these stout and to 27 cm. long); staminodes lacking in 
pistillate flowers; stems robust, erect to scandent in mature plants, 
with the internodes usually elongate. 

Pinnae rhombic-lanceolate and strongly sigmoid, 6-9 on each side of 
a pale-backed rachis; flowers drying black in both sexes. 

C. oblongata 

Pinnae linear to lanceolate, not sigmoid but sometimes falcate toward 
the apex, generally more than 10 on each side of a green-backed 
rachis; flowers not drying black. 



222 FIELDIANA: BOTANY, VOLUME 24 

Branches of the inflorescence short, stiff, spinose-tipped, promi- 
nently ridged and angled C. aequalis 

Branches of the inflorescence not ridged, angled, and pointed. 
Pinnae narrowly linear, about 30 on each side of the rachis; 

plants colonial C. Schippii 

Pinnae lanceolate; plants solitary or cespitose. 
Inflorescence breaking through the long sheaths on a sprawl- 
ing, clambering, or scandent stem, subtended by 3-4 
spathes, the uppermost spathe incomplete and often the 
only one apparent; leaves variable but at maturity to 3 
meters long with reflexed pinnae narrowed to a very 

slender callose-indurate base C. elatior 

Inflorescences erect from the orifices of the sheaths, subtended 

by about 6 spathes. 

Pinnae with the midnerve prominent above, yellow and 
shining below, the remaining nerves less conspicuous; 
pistillate spadix elongate with 18-25 branches only 

6-9 cm. long C. carchensis 

Pinnae with the midnerve and 2 secondary nerves on each 
side prominent, yellowish and shining below; pistillate 
inflorescence with few (3-6) elongate branches. 

C. vulgata 

Inflorescences of both sexes spicate or with few (rarely as many as 10) 

branches; staminodes present in pistillate flowers (except C. quet- 

zalteca?) ; stems slender and generally low, usually densely annulate. 

Inflorescences breaking through the persistent sheaths at the nodes 

(see also C. digitata). 

Peduncles of the inflorescence short, scarcely as long as the few 
short branches. 

Leaves pinnate; plants robust, cespitose C. erumpens 

Leaves simple; plants slender, low and colonial . . .C. brachypoda 
Peduncles of the inflorescence elongate; plants solitary. 

Pistillate inflorescence spicate; staminate with 2-4 very densely 
flowered pendulous branches C. adscendens 

Pistillate inflorescence with 4-10 branches; staminate with 8-10 
branches. 

Inflorescences of both sexes with about 10 branches; central 
pinnae to 20 cm. long, 3.5 cm. wide C. pulchra 

Pistillate inflorescence with 4 branches; staminate with 8 
branches; central pinnae to 15 cm. long, 2 cm. wide. 

C. partrisecta 

Inflorescences erect, emerging through the orifices of the sheath or 
from the base of the stem. 

Leaves rounded and undivided at the apex C. Tuerckheimii 

Leaves cleft at the apex. 

Petiole densely rugose; leaves entire or pinnate, then with pinnae 

prominently auriculate at the base C. tenerrima 

Petiole smooth or only slightly verrucose above; pinnae not 

prominently auriculate. 

Leaves simple; inflorescence very long-pedunculate (80-100 
cm.), subtended by 9-10 closely sheathing spathes. 

C. stricta 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 223 

Leaves pinnate; inflorescence less than 30 cm. long with 7 or 

fewer spathes. 
Leaf sheaths elongate, cylindric, closely sheathing the 

caudex. 
Pinnae about 16 on each side, falcate, prominently 3-5- 

nerved C. quetzalteca 

Pinnae about 7 on each side, more or less sigmoid, only 

the central nerve prominent C. digitaia 

Leaf sheaths short, open, imbricate. 

Pinnae markedly sigmoid, short and broad (generally 4 
times as long as broad or less), about 10 on each 

side of the rachis C. Pachecoana 

Pinnae lanceolate-attenuate or falcate-lanceolate, gen- 
erally about 5 times as long as broad. 
Fruit oblong-ellipsoid; pinnae 10-14 on each side, 

prominently 3-nerved C. stenocarpa 

Fruit globose or ovoid; pinnae about 8 on each side, 
the central nerve more prominent than the others. 

C. Lehmannii 



Chamaedorea adscendens (Dammer) Burret, Notizbl. Bot. 
Gart. Berlin 11: 737. 1933; 11: 1039. 1934. Kinetostigma adscendens 
Dammer, Notizbl. Bot. Gart. Berlin 4: 172. 1905. Tuerckheimia 
adscendens Dammer ex Donn. Smith, Enum. PI. Guat. 7: 53. 1905, 
nomen. 

Moist or wet, mixed forest, often in rocky places, 700 meters or 
lower; Alta Verapaz (type from summit of Cerro Hacoc, near Cubil- 
giiitz, Tuerckheim 8770; collected at other localities in the same 
general region); Izabal. British Honduras. 

Plants slender, 2.5 meters high or lower, sometimes flowering when acaulescent 
or nearly so, the caudex commonly about 1 cm. thick; leaves few, generally about 
6, dull green with a velvety appearance, the cylindric sheath 6 cm. long or shorter, 
the petiole 7 cm. long or more, the rachis short; leaves variously and often unevenly 
divided into 1-6 segments on each side, the longest 16 cm. long and 2.5 cm. wide 
with a prominent but scarcely elevated midnerve pale below and numerous incon- 
spicuous nerves of lesser orders, or the terminal pair often much broader and 2-9- 
nerved, or sometimes one side undivided; inflorescences solitary and penetrating 
the sheaths at the nodes, then divaricate-ascending to erect, the staminate peduncle 
slender, as much as 40 cm. long, the spathes 8-9, slender, the rachis very short, the 
branches 2-5, slender, as much as 15 cm. long and pendulous with cream-colored 
nerveless flowers 2.5 mm. long superficial on the branchlets in a densely spiralled 
arrangement, the calyx pale, shallowly 3-lobed, the petals briefly connate at the 
base, distinct and not united above, the anthers apically entire and longer than 
the filaments, pistillode columnar; pistillate spadix long-pedunculate, simple, the 
floriferous part about 10 cm. long, slender, the nerveless flowers remote and scarcely 
sunken, the calyx annular, briefly 3-lobed, about 1 mm. high, petals very shortly 



224 FIELDIANA: BOTANY, VOLUME 24 

connate basally, broadly imbricate above, acute, staminodes present, abortive car- 
pels adherent to the corolla in fruit; fruit black at maturity, globose or ovoid- 
globose, 8 mm. long; seed 6 mm. long. 

A specimen of this species (Schipp 1223, Gray Herbarium) is 
anomalous in having sterile (?) pistillate flowers among the predom- 
inant staminate ones in a typical staminate spadix. Staminate 
spadices at maturity are very densely flowered, superficially resem- 
bling such species as C. Arenbergiana or C. Tepejilote but in bud the 
flowers are clearly distinct. 

Chamaedorea aequalis Standl. & Steyerm. Field Mus. Bot. 23: 
196. 1947. 

Moist or wet, mixed forest, sometimes growing with Liquidam- 
bar, 1600-2000 meters; endemic; Huehuetenango (type from Cerro 
Negro, 2 miles west of Las Palmas, Sierra de los Cuchumatanes, 
Steyermark 51666); Solola (southern slopes of Volcan de Atitlan); 
Quezaltenango (slopes of Volcan Zunil). 

Caudex erect, 3-4.5 meters high, 1.5-2 cm. thick, the upper internodes 11-15 
cm. long; leaf sheaths open and oblique, about 9 cm. long, prominently costate, 
the petiole to 19 cm. long, 3 mm. thick; leaf blades 37 cm. long and 23 cm. wide or 
larger; pinnae 17 or more on each side, thin, concolorous, regularly disposed, some- 
what lustrous, the apical ones not wider than the others, connate at the base, the 
pinnae all narrowly lanceolate, not sigmoid, 13-30 cm. long, 1.5-4 cm. wide, nar- 
rowly and equally attenuate-acuminate, gradually narrowed to the very narrow 
base, mostly 3-nerved, the nerves not keeled above, the secondary nerves numer- 
ous, almost as stout as the primary ones; peduncles ascending or suberect from 
below the leaves, slender, 21-30 cm. long, scarcely more than 2 mm. thick above; 
spathes 6, tubular, imbricate, the lower ones very short, the middle ones about 
12 cm. long, 5 mm. in diameter, appressed ; staminate spadix with about 25 branches 
from an axis 9.5 cm. long, the branches spreading, sharply angled and ridged, to 
12 cm. long, the lower ones again once forked or branched, the nerveless flowers 
superficial, about 3.5 mm. high, the calyx pale, shallowly 3-lobed, about 1 mm. 
high, petals valvate, not connate at the tips, the stamens with prominent filaments 
and anthers entire at the apex, pistillode nearly cylindric, rounded at the tip; pistil- 
late spadix 10-13 cm. long and broad, the branches about 10, sharply angled and 
ridged, erect-patent, the lowest ones furcate or branched, the rest simple, 9-10.5 
cm. long, slender, red in fruit, the flowers scarcely immersed, the fruits remote, 
spirally arranged; corolla explanate below the fruit, the petals equal, broadly ovate, 

2 mm. long, nerveless, subapiculate, the calyx scarcely more than 2 mm. broad, the 

3 lobes broadly rounded; staminodes lacking; fruit globose, 6 mm. in diameter when 
dry; seed almost as long as the fruit, globose, pale grayish brown. 

This species is unusual among Guatemalan representatives of the 
genus in the clearly ridged branchlets of the inflorescence. In this 
respect it resembles C. elegans but is probably even more closely re- 
lated to C. Liebmannii of Mexico. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 225 

Chamaedorea Aguilariana Standl. & Steyerm. Field Mus. Bot. 
23: 196. 1947. Chilak (Huehuetenango) ; Pacaya; Molinillo. 

Moist or wet, mixed, mountain forest, 600-2000 meters; endemic; 
Santa Rosa; Escuintla; Suchitepquez; Chimaltenango; Quezalte- 
nango (type collected in quebrada along the old road between Finca 
Pirineos and Patzulin, Standley 86890 pro parte); San Marcos; 
Huehuetenango. 

Plants solitary, the caudex slender, erect, green, 1-3 meters high, 8 mm. thick, 
conspicuously annulate, the internodes mostly 1.5-2 cm. long; leaf sheaths 9-10 cm. 
long, 1-1.5 cm. broad, oblique at the apex; leaf blades oblong in outline, mostly 
30-40 cm. long, the rachis very slender; 4-5 pinnae on each side, regularly distrib- 
uted, alternate, the apical ones confluent, 18-20 cm. long and 2.5-5 cm. wide, thin, 
sigmoid, falcate-acuminate, slightly paler beneath, central and submarginal nerves 
prominent and very slightly keeled above, pale and shining below as are the 2-4 
slender secondary nerves on each side; inflorescences borne among the leaves, sub- 
tended by 3-4 tubular, thin spathes 6-7 mm. in diameter, the peduncles 8-22 cm. 
long; staminate spadix with 11-14 slender pendulous branches about 12 cm. long 
from a short axis, floriferous for their whole length, the green, essentially super- 
ficial flowers strongly nerved when dry, 3.5 mm. high, rather densely spiralled, the 
calyx 0.75 mm. high, rather shallowly and not imbricately 3-lobed, the petals very 
shortly connate in a compressed linear base, expanded above and connate at their 
tips, the corolla opening by lengthwise slits, stamens with short filaments and ob- 
long anthers entire at the apex, the pistillode 3-angled and truncate; pistillate 
spadix with 6-18 branches 6-10 cm. long from a short to moderate rachis, the 
green flowers 2 mm. high, strongly nerved when dry, loosely spiralled and essen- 
tially superficial, the calyx 1 mm. high, 3-lobed; petals imbricate, staminodes lack- 
ing; pistil ovoid, with sessile stigmas; fruit orange, perhaps darkening at full 
maturity, globose, 6 mm. in diameter. 

Differences between C. Aguilariana and C. concolor of Mexico are 
few if any, and it is possible that further study will show them to be 
identical. The probable relationships to plants of this complex from 
Alta Verapaz are discussed with C. micrantha. 

The name "molinillo" given to this and other species relates to 
the fact that the base of the stem with the cluster of stiff roots is 
used to stir coffee and other hot liquids. The species was named for 
Don Jos Aguilar, formerly Director of the Finca Nacional La Aurora. 

Chamaedorea Arenbergiana Wendl. Index Palm. 66. 1854; 
Kerchove, Les Palmiers 75. /. 33. 1878; Bot. Mag. pi. 6838. 1885. 
Spathoscaphe Arenbergiana Oerst. Vid. Medd. Kjoebenhavn 1858: 
30. 1859; AmeY. Centr. 15. pi. 7, f. 29-37. 1859. Nunnezharoa Aren- 
bergiana Kuntze, Rev. Gen. 2: 730. 1891. Chim (San Marcos). Fig- 
ure 41. 







FIG. 41. A, Chamaedorea elegans. B, C. Ernesti-Augustii (m, XI). C, 
C. Tepejilote. D, C. Arenbergiana (k, calyx in bud; m, X 1^). a-gr, staminate; 
h-n, pistillate: a, portion of rachilla (X 2%); b, flower (X 6); c, corolla (X 6); 
d, calyx ( X 6) ; e, flower expanded ( X 6) ; /, pistillode ( X 8) ; g, stamens in front, 
back, and side views (X 8); h, portion of rachilla (X 2%); i, flower (X 6); j, 
corolla (X 6) ; k, calyx (X 6) ; I, pistil (X 6) ; m, fruit (X 2) ; n, perianth in fruit 
(X2). 

226 







Hm H. 



FIG. 41 (continued). E, Chamaedorea Rojasiana. F, C. geonomaeformis. G, 
C. oblongata. H, C. Pachecoana. a-g, staminate; h-n, pistillate: a, portion of 
rachilla (X 2%); b, flower (X 6); c, corolla (X 6); d, calyx (X 6); e, flower ex- 
panded (X 6) ; /, pistillode (X 8) ; g, stamens in front, back, and side views ( X 8) ; 
h, portion of rachilla (X 2%); i, flower (X 6); ;, corolla (X6); k, calyx (X 6); 
/, pistil (X 6); m, fruit (X 2); n, perianth in fruit (X 2). 

227 



228 FIELDIANA: BOTANY, VOLUME 24 

Moist or wet, mixed forest, 300-1500 meters; described from cul- 
tivated plants believed to be of Guatemalan origin; Alta Verapaz; 
Izabal; San Marcos; Quezaltenango; Huehuetenango. Honduras to 
Panama. Represented in Herbarium of Chicago Natural History 
Museum by three type photographs (Negatives 20754, 2106, 25382). 

Plants 4.5 meters high or lower, sometimes flowering while still acaulescent, 
the stems green, 2-3 cm. in diameter; leaves dark green, erect-spreading, 1-2.5 
meters long, the sheath obliquely open at the apex, sometimes 40-45 cm. long, the 
petioles 35-45 cm. long; leaf segments 8-10 on each side of a pale-backed rachis, 
rather remote, with 5-8 prominent primary nerves keeled above, shining and pale 
below and numerous nerves of lesser orders, elongate-oblong or oblong-lanceolate, 
often somewhat falcate, very long-acuminate and drooping at the tips, the largest 
about 60 cm. long and 8-10 cm. wide, most of the segments smaller or at least much 
narrower, green on both surfaces; inflorescences arising beneath the leaves, sub- 
tended by 5 spathes, the staminate spadices 30-40 cm. long with 8-10 pendulous 
branches 12-15 cm. long, the brownish flowers crowded on the axis in 6 spiralling 
rows, the calyx gamosepalous, longer than the corolla and enclosing it in bud, about 
half as long at anthesis, shallowly 3-lobed, not prominently nerved, the corolla 
united with the stamen filaments and pistillode in a very short basal stipe but dis- 
tinct with spreading valvate lobes 2.5 mm. long above, the stamens with prominent 
distinct filaments and anthers divaricate basally, pistillode columnar, apically 
3-lobed, equaling the stamens; pistillate spadix usually simple, sometimes furcate, 
about 30 cm. long, the peduncle very thick, erect, much longer than the fruiting 
portion of the spadix, this usually 5-12 cm. long and about 1.5 cm. thick or broader, 
flowers very densely crowded in perhaps as many as 20 rows and somewhat sunken 
in the axis, the sepals connate in a shallowly 3-lobed membranous cup nearly as 
high as the broadly imbricate, scarcely nerved, shortly pointed petals, staminodes 
absent, pistil ovoid, 3-lobed with 3 short, recurved, sessile stigmas, the abortive 
carpels usually separating from the fruit and adhering to the perianth; fruits 
crowded, subglobose or transverse-oblong, about 12 mm. long and sometimes 
19 mm. broad; seed oblong, 10-12 mm. long, 9-10 mm. broad. 

Pistillate inflorescences of this species are generally simple or fur- 
cate, differing in this respect from C. Casperiana, with which it may 
eventually prove conspecific. 

Chamaedorea brachypoda Standl. & Steyerm. Field Mus. Bot. 
23: 198. 1947. 

Dense, wet, mixed, lowland forest, 150 meters or lower; endemic; 
Izabal (type collected between Bananera and La Presa, base of Sierra 
del Mico, Steyermark 39185; collected also between Virginia and Lago 
de Izabal and in the vicinity of Quirigua) . 

Plants rhizomatous and colonial (?), the caudex slender, pale green, about 1 
meter high, 6-7 mm. thick, the internodes 6-7.5 cm. long; leaf sheaths about 6 cm. 
long, tubular, 8 mm. broad, striate-nerved, oblique at the apex, more or less per- 
sistent, the slender petiole 9-10 cm. long; leaf blades simple, about 25 cm. long 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 229 

and wide, cuneate-acute at the base, the rachis about 12 cm. long, pale below, the 
primary nerves 11-13 on each side, prominent and keeled above, the secondary 
nerves much more slender than the primary ones, the blade deeply bilobate at the 
apex, the lobes acute or short-acuminate, divergent at an angle of 90 degrees or 
more, about 16 cm. long and 10 cm. wide, almost concolorous; inflorescences break- 
ing through persistent old sheaths at the node below the living leaves, the peduncle 
3-5 cm. long, slender but stiff, ascending; spathes 6, very short, the lowest scarcely 
more than 8 mm. long, the uppermost as much as 2.5 cm. long, 4 mm. broad, acute 
or acuminate at the apex; branches of the staminate spadix 5-6, very slender, 
9-10.5 cm. long, laxly flowered, the yellow flowers about 3.5 mm. high, scarcely 
immersed, the calyx 1 mm. high, shallowly 3-lobed, petals united for about 1 mm. 
at the base and adnate to the stamen filaments and pistillode in a short stipe, free, 
fleshy, valvate above and somewhat inflexed but not connate at the apex, the sta- 
mens with very short filaments, anthers subbasally attached and bifid apically for 
more than half their length, the pistillode sharply 3-angled, ventricose at or above 
the middle, narrowed to a truncate angled cap; pistillate spadix with 6-8 branches 
about 6 cm. long, the flowers in young bud slightly immersed, depressed-globose, 
with shallowly 3-lobed calyx; petals imbricate at the base, the apex deltoid and 
probably deciduous in fruit, staminodes present; immature fruit ellipsoid. 

This species, if cultivated material corresponding with it is typ- 
ical, is probably colonial with long slender underground rhizomes. 

Chamaedorea carchensis Standl. & Steyerm. Field Mus. Bot. 
23: 199. 1947. 

Known only from the type locality, wet rocky forest along banks 
of Rio Carcha, between Coban and San Pedro Carcha, Alta Verapaz, 
1350 meters, Standley 90160. 

Plants acaulescent; leaves large, the sheaths indurate, 12 cm. long, almost 2 cm. 
thick, narrowed to the petiole; petiole 1 meter long or slightly longer, narrowly 
sulcate, almost 1 cm. thick; pinnae about 15 on each side, subequal, regularly in- 
serted, alternate, linear-lanceolate, the terminal ones scarcely wider or in young 
leaves twice as wide as the lower pinnae, the middle pinnae about 40 cm. long and 
3.5 cm. wide, straight, not sigmoid, equally attenuate-acuminate, gradually nar- 
rowed to the base, the base itself 10-15 mm. wide, slightly indurate; pinnae con- 
spicuously 1-nerved and with 2 more slender submarginal nerves, the secondary 
nerves very slender and inconspicuous; pistillate inflorescences arising from the 
very base of the plant, erect, the peduncles 34-50 cm. long, rather stout, 4 mm. 
thick near the apex; spathes 6, imbricate, appressed, 8-10 mm. broad, the lowest 
only 3.5 cm. long, the uppermost about 12 cm. long, oblique at the apex, obtuse 
and submucronate, densely striate-nerved; branched portion of the pistillate spadix 
20-26 cm. long, 7-8 cm. broad, the rachis rather stout, 14-18 cm. long, the branches 
18-25 and 6-9 cm. long, stout, green, arcuate-erect, sparsely flowered; flowers 
scarcely immersed, the alveolae 1-1.2 mm. long, orbicular or broadly oval; calyx 
2 mm. broad, thickened in age, deeply 3-lobate, the lobes rounded-ovate, very 
obtuse; petals 3 mm. long, rounded-ovate, obtuse but apiculate, thickened in age; 
young fruit scarcely 5 mm. long, green, ovoid or broadly oblong but probably 
nearly globose in fruit, probably maturing black or glaucous. 



230 FIELDIANA: BOTANY, VOLUME 24 

The plant may be rare, since we have found it in only the one 
locality. The very young plants have small, simple, deeply bifid 
leaves as in C. elatior, to which it appears related. The type locality, 
the wooded rocky banks of Rio Carcha, is one of the few places left 
close to Coban that retain something of the original, highly varied 
forest that presumably covered formerly a great part of the Coban 
region. Most such forest, if on land that could by any possible means 
be cultivated, long ago was cleared for agricultural purposes. 

Chamaedorea Casperiana Klotzsch in Otto & Dietr. Allg. 
Gartenz. 20: 363. 1852. Stephanostachys Casperiana Oerst. Vid. 
Medd. Kjoebenhavn 1858: 27. 1859; Amer. Centr. 15. pi 7, f. 1-11. 
1863. Nunnezharoa Casperiana Kuntze, Rev. Gen. 2: 730. 1891. 

Described from cultivated plants, said to have been collected in 
Guatemala by Warscewicz; represented in Herbarium of Chicago 
Natural History Museum by three photographs (Negatives 20753, 
25385, 29891) of cultivated material distributed by Wendland and 
agreeing with Klotzsch's descriptions. 

Plants solitary with stems to 0.5 meters high (or probably more), thickened to 
5.5 cm. in diameter basally, about 2 cm. in diameter above; leaves erect-spreading, 
about 1 meter long, the sheath about 20 cm. long, clasping the stem and opening 
obliquely at the apex, the petiole about 27.5 cm. long, pale below as is the rachis, 
the pinnae 6-7 on each side of the rachis, 5-10-nerved, elongate-lanceolate, falcate, 
with pendulous long-acuminate tips, the lower pinnae reflexed-spreading, the upper 
somewhat broader than the remainder, the median (and longest) pinnae 32.5- 
37.5 cm. long, 5-6 cm. wide at the middle, 1.8 cm. wide at the base; spadices borne 
below the leaves, subtended by 4 somewhat inflated spathes, sometimes an incom- 
plete fifth spathe borne near the top of the peduncle, the staminate spadices of 7-10 
or more pendulous branches to 12 cm. long, densely crowded with white flowers in 
about 6 rows, the calyx probably membranous, gamopetalous, and conspicuously 
cupular in bud (from photos), the corolla campanulate with spreading valvate- 
deltoid lobes; the pistillate spadices erect with about 8 stiff erect branches from a 
very short axis, the branches to 7.5 cm. long, densely crowded with flowers in about 
6 rows, the calyx gamosepalous, membranous and conspicuously cupular in bud 
(from photos), the petals broadly imbricate; fruit not described. 

The species has not appeared in recent collections from Guate- 
mala and its provenance remains uncertain. It is very similar to 
C. Arenbergiana, differing in its smaller leaves and in the more numer- 
ous branches of the pistillate spadix. 

Chamaedorea digitata Standl. & Steyerm. Field Mus. Bot. 
23: 200. 1947. 

Moist or wet, mixed, mountain forest, 1400-2600 meters; en- 
demic; El Progreso (Sierra de las Minas); Huehuetenango (type 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 231 

collected in Liquidambar forest, Cerro Negro, 2 miles east of Las 
Palmas, Sierra de los Cuchumatanes, Steyermark 51683). 

Caudex slender, erect, green, to 1 meter high or more, 5-7 mm. thick, the inter- 
nodes unequal, short or elongate; leaves rather small, the sheath 16-21 cm. long, 
tubular, oblique at the apex, 6-13 mm. thick, conspicuously costate-nerved, the 
sheaths sometimes (the lowest ones) only 8 cm. long, the petioles slender, 14-17 cm. 
long; leaf blades generally 17-28 cm. long, the pinnae 4-6 on each side, regularly 
inserted, thin, slightly paler beneath, lanceolate or broadly lanceolate, sigmoid, 
6-13 cm. long, 2.5-3 cm. wide, narrowly attenuate-acuminate, narrowed to the 
base, 4-6-nerved, the primary nerve central, pale below and slightly elevated 
above, with 3-4 less conspicuous lateral nerves and numerous finer nerves; in- 
florescences radical in acaulescent plants and apparently breaking through the 
sheaths at the nodes, the peduncles very slender, about 18 cm. long, 1.5 mm. thick, 
spathes about 6, closely sheathing, tubular, brownish, striate-nerved, 3-4 mm. in 
diameter, subacute at the apex; rachis of the staminate spadix only 5-6 mm. long, 
the 4 branches slender, remotely flowered, 7.5-9 cm. long, the flowers depressed- 
globose, slightly immersed, the calyx 1.7 mm. broad, very shallowly lobate, the 
lobes truncate; petals ecostate, rounded-ovate, valvate, obtuse; pistillate spadix 
with 3-5 slender erect green branches 2-3.5 cm. long, the flowers remote, shallowly 
immersed, calyx very shallowly 3-lobate, 2.5 mm. broad, petals ovate-rounded, 
often broader than long, slightly more than 2 mm. long, somewhat 3-nerved out- 
side; immature fruits 5-6 mm. long, subglobose or ovoid-globose, rounded at the 
base and apex; staminodes present. 

Specimens originally assigned to this species are somewhat vari- 
able. The type and a staminate plant from El Progreso have inflo- 
rescences from the base of a short stem and elongate petioles. Others 
from Solola and San Marcos have axillary inflorescences and short 
petioles only 3-6 cm. long. These appear to be more closely related 
to C. Pachecoana. The description, therefore, is based principally 
on the type. 

Chamaedorea elatior Martius, Linnaea 5: 205. 1830. Nunne- 
zharoa elatior Kuntze, Rev. Gen. 2: 730. 1891. C. scandens Liebm. 
in Martius, Hist. Nat. Palm. 3: 308. 1849. C. resinifera Wendl. in 
Otto & Dietr. Allg. Gartenz. 21: 179. 1853. Nunnezharoa resinifera 
Kuntze, Rev. Gen. 2: 731. 1891. C. desmoncoides Wendl. in Otto & 
Dietr. Allg. Gartenz. 21: 177. 1853. Nunnezharoa desmoncoides 
Kuntze, Rev. Gen. 2: 720. 1891. Chiang (San Marcos). 

Moist or wet, mountain forest, 1100-1500 meters; Alta Verapaz 
(near Secoyocte", Finca Sepacuite); San Marcos (Volcan de Taju- 
mulco); Huehuetenango (region of Barillas). Eastern and southern 
Mexico. 

Stems sprawling or subscandent, often with arching stems, but when young 
nearly erect, the caudex 1.5-2 cm. in diameter or less; leaves with elongate tubular 



232 FIELDIANA: BOTANY, VOLUME 24 

sheaths often covering two nodes and to about 30 cm. long, clothing much of the 
stem, the blades long-persistent; petioles very short (mostly on adult leaves) to 
elongate; blades very long, sometimes 3 meters or more in robust adult individuals, 
entire and deeply bifid when juvenile, progressively more divided up the stem with 
numerous linear-lanceolate deep green pinnae becoming almost equally attenuate 
at each end in adult leaves and then strongly indurate-calloused at the very narrow 
base, and reflexed, median adult pinnae about 35 cm. long or less and to 3 cm. wide, 
not at all sigmoid, with a prominent central nerve and numerous but inconspicuous 
secondary and tertiary nerves; inflorescences breaking through the sheaths, sub- 
tended by 3-4 spathes, only the upper (and often incomplete) one generally visible, 
the peduncle usually recurved; staminate spadix 20 cm. long and broad or larger, 
the branches numerous, slender, naked near the base, rather laxly flowered above, 
divaricate at a right angle or somewhat reflexed; pistillate spadix large, the 
branched portion to 40 cm. long and 30 cm. broad, with rather few to very many 
simple, remotely flowered branches as much as 30 cm. long, these pale orange or 
salmon-colored or dull brownish in fruit, the flowers not sunken in the axis, the 
calyx deeply lobed, petals imbricate, rounded and shortly apiculate, staminodes 
lacking; fruit globose and black with a glaucous bloom at maturity, green when 
immature, about 10 mm. in diameter. 

The Guatemalan specimens referred here correspond well with 
Chamaedorea desmoncoides in their very short petioles and profusely 
branched inflorescences. Chamaedorea elatior, however, is variable, 
not only in the various stages through which the leaves go as they 
develop along the stem, but in size of parts, even in the same region. 
Thus, elongate or very short petioles may be found on the same plant, 
depending on the position of the leaf. The number of pinnae and of 
branches of the pistillate spadix appears to vary sufficiently to make 
the separation of forms cited in synonymy not only difficult but 
illogical. 

This species is distinct in habit. The rather stout stems arch and 
either sprawl over the ground or are supported on other vegetation. 
The epithet desmoncoides would be most appropriate were it the old- 
est available since plants are readily mistaken for Desmoncus when 
not examined closely. 

Chamaedorea elegans Martius, Linnaea 5: 204. 1830. Collinia 
elegans Liebm. ex Oersted, Vid. Medd. Kjoebenhavn 1858: 6. 1859. 
Nunnezharoa elegans Kuntze, Rev. Gen. 2: 730. 1891. Neanthe bella 
0. F. Cook, Science 86: 120. 1937, nomen without Latin description; 
Nat. Hort. Mag. 17: 1-12, /. 1-8. 1938, without Latin description. 
Pacaya; Pacayito; Kiik (Quecchi). Figure 41. 

Moist or wet, usually dense, mixed, mountain forest, 1400 meters 
or lower; Pete"n; Alta Verapaz. Eastern and southern Mexico. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 233 

Plants very slender, in age as much as 2 meters high or even taller, but often 
flowering when very small and not more than 30 cm. high, then essentially acau- 
lescent, the caudex erect or decumbent, 8-16 mm. in diameter, green, densely 
annulate with short internodes; leaves few and small, the sheath long, slender, 
open nearly to the base, the petiole slender, 12-27 cm. long, the rachis very slender 
and pale-backed; pinnae 11-20 on each side of the rachis, linear to narrowly lanceo- 
late, long-attenuate, 12-20 cm. long, 1-2 cm. wide, the primary nerve central, pale, 
prominent, slightly elevated above but not keeled, secondary nerves generally less 
prominent, 2 on each side, evenly spaced, tertiary nerves numerous and fine; in- 
florescences erect from the sheaths, the peduncle 10-13 cm. long or more, subtended 
by 4-7 (o") or 6-9 ( 9 ) tubular sheaths split at the apex, the spadices with few or 
numerous, simple or branched green rachillae with sharp ridges, the pistillate rachil- 
lae becoming orange in fruit; flowers sessile, remotely spiralled, the staminate 2 mm. 
high, pale yellow, prominently nerved when dry, the calyx 0.75 mm. high, shal- 
lowly 3-lobed; petals connate nearly to their tips, the apex of the corolla with a 
3-angled opening; stamens with short filaments and entire anthers, these scarcely 
visible beneath the flaring 6-angled apex of the pistillode; pistillate flowers 2.5 mm. 
high, nerved when dry, the calyx 1 mm. high, rather deeply 3-lobed, petals connate 
except for a 3-angled aperture at the tip, staminodes lacking, ovary depressed- 
globose with sessile stigmas; fruit globose, black at maturity, about 6 mm. in 
diameter. 

Chamaedorea elegans has been separated as the single species in 
Collinia by Burret. Despite the unusual union of perianth parts, 
the species is obviously closely related to others, such as C. Lieb- 
mannii and C. humilis of Mexico, in which the corolla is lightly imbri- 
cate in the pistillate flowers; fruiting specimens are distinguished by 
the valvate lobes of the corolla, which splits evenly into three parts. 

The Guatemalan specimens referred here have been considered 
distinct by 0. F. Cook, who used the invalid name Neanthe bella for 
them. When plants from the entire range of the species are consid- 
ered, it is difficult to find differences that are of specific significance. 
In general, however, plants from Guatemala are more delicate than 
those from the northern part of the range and they rarely have other 
than simple branches in the inflorescence of both sexes. 

Chamaedorea Ernesti-Augustii Wendl. in Otto & Dietr. Allg. 
Gartenz. 20: 73. 1852; Index Palm. 12: 58. 1854; Hook. f. in Bot. 
Mag. pi 4831, 4837. 1855; Flore des Serres ser. 2, 3: 133. pi 1357. 
1858. Eleutheropetalum Ernesti-Augustii Oerst. Vid. Medd. Kjoeben- 
havn 1858: 7. 1859. Nunnezharoa Ernesti-Augustii Kuntze, Rev. 
Gen. 2: 730. 1891. Figure 41. 

Dense, wet forests, often on limestone, mostly at low elevations 
but to 1100 meters in Alta Verapaz; Alta Verapaz; Huehuetenango; 
Izabal; Pet&i. Southern Mexico to Honduras, described from culti- 
vated plants supposedly from Tabasco, Mexico, and represented in 



234 FIELDIANA: BOTANY, VOLUME 24 

the Herbarium of Chicago Natural History Museum by type photo- 
graphs (Negatives 20806, 20806a, 25388). 

Plants to 2 meters high or more, the slender stems to 13 mm. in diameter, 
sometimes flowering while nearly stemless; leaves spreading, simple, broadly cune- 
ate-obovate in outline, deeply cleft at the apex, dull dark green above, dull green 
below, the sheath obliquely open above the middle, 8-10 cm. long, the petiole 
8-20 cm. long, pale-centered below as is the rachis, this 17-28 cm. long, the blade 
22-25 cm. long on the upper margin, 20-25 cm. wide at the tip of the rachis, the 
subscarious margins shortly emarginate at the tips of the 13-16 nerves on each 
side, toothed between the nerves, the latter pale, dull, and scarcely prominent be- 
low, prominently keeled above; inflorescences solitary from orifices of the sheaths, 
but sometimes persisting below the leaves in fruit, subtended by 4 ( d" )-5 ( 9 ) 
closely sheathing tubular spathes, the uppermost of which is slightly shorter than 
to exceeding the erect peduncle, the staminate spadices with an elongate peduncle 
to 30 cm. long or more, copiously branched with slender, green, divaricate to pendu- 
lous, simple branches to 13 cm. long from a rachis to 15 cm. long, the flowers 2.5 
mm. high, seated in shallow depressions in a loose spiral, the pale yellow, mem- 
branous, nerveless calyx about 1 mm. high, rather deeply 3-lobed and brown- 
margined, the bright orange corolla with petals connate in a shortly stipitate base, 
free, fleshy, nerveless and valvate above, the stamens with filaments shorter than 
the anthers, these deeply bifid at base and apex, pistillode columnar, expanded into 
a truncate 6-angled cap; the erect pistillate spadices with peduncles to 70 cm. long, 
the rachis to 27 cm. long, simple, thick, dull green at anthesis, becoming orange or 
vermilion and thicker in fruit, the flowers about 3.5 mm. high, separated in promi- 
nent elliptic depressions in a loose spiral, the yellowish calyx of 3 lightly imbricate 
nerveless sepals 1.5 mm. high, the bright orange petals thin, slightly nerved and 
very narrowly imbricate at the base in bud though at anthesis appearing valvate 
throughout, above becoming fleshy, nerveless, and cucullate, the cucullate portion 
usually deciduous as fruit develops, staminodes present, prominent, pistil of 3 
nearly free carpels lightly connate centrally near the base at anthesis, stigmas 
sessile and recurved, the abortive carpels adherent to the perianth in fruit; fruit 
subglobose to generally ellipsoid, green to blue-green becoming black at maturity, 
to 14 mm. long, 8 mm. in diameter with remains of a single stigma at the base and 
pale depressions where abortive carpels have torn away, the seed to 11 mm. long, 
7 mm. in diameter. 

Chamaedorea Ernesti-Augustii is frequently confused with C. ge- 
onomaeformis, from which it differs in obvious floral characters and 
in the keeled nerves of the broader leaves. A closely related but 
stouter species, C. Sartori, differs principally in its leaves with several 
sigmoid pinnae on each side and 4-6 branches of the pistillate spadix. 
The last ranges from eastern Mexico to Honduras and may be ex- 
pected in Guatemala, though it has not been recorded. 

Chamaedorea erumpens H. E. Moore, Gentes Herb. 8: 232. 
1951. 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 235 

Dense, wet, lowland forests at altitudes of 500 meters or less; 
described from plants supposedly from British Honduras; Pete'n 
(Tikal district). 

Plants cespitose in clumps of as many as 40 slender green stems to 3 meters 
high or more, 11-17 mm. in diameter, the internodes elongate; leaves about 4, pin- 
nate, the cylindric sheaths 16.5-20 cm. long, striate-nerved, the petiole 4.5-8 cm. 
long, the rachis green below, 18-47.5 cm. long with 5-15 deep green pinnae on each 
side, the terminal pair sometimes scarcely larger than those below, sometimes 
broader and then to 9 cm. wide and 7-9-nerved and lower pinnae 4-5 on each 
side of a foreshortened rachis; pinnae lanceolate, tapered subfalcately below to a 
short-acuminate or acute pendulous tip, the median largest, to 27.5 cm. long, 
1.6-3 cm. wide, all but the terminal pair with a central primary nerve prominent 
below but scarcely elevated above and a pair of lateral and submarginal nerves 
on each side, the remainder of two orders, numerous and inconspicuous; inflores- 
cences solitary, erumpent through the sheaths at the nodes or persisting below the 
leaves, the peduncles short, 4-7 cm. long, subtended by 5 short, closely sheathing 
spathes, the uppermost sometimes exceeding the peduncle and split along the lower 
side, the green staminate spadix with 6-11 erect or spreading, simple or rarely fur- 
cate branches 7.6-15 cm. long from a rachis 2-4.5 cm. long, the flowers yellow, 
somewhat sunken in a loose spiral, the nerveless sepals very shortly connate at the 
base, imbricate above, 1 mm. high, the petals nerveless, valvate above a shortly 
connate base, fleshy, not connate at the tip, the stamens with short filaments and 
anthers bifid at the tip, pistillode columnar, 3-lobed at the apex; pistillate spadix 
with 4-6 simple erect deep green branches 9-10.5 cm. long from a rachis 1-2.5 cm. 
long, becoming thickened and orange in fruit, the yellow flowers somewhat sunken 
in a loose spiral, sepals green drying yellow, imbricate, 1-2 mm. high, petals imbri- 
cate, nerveless, acutish, 3 mm. high, drying brown and slightly nerved, staminodes 
present, pistil subglobose with sessile stigmas; fruit black at maturity, globose, 
about 8 mm. in diameter. 

Chamaedorea falcifera H. E. Moore, Principes 2: 68, fig. 45. 
1958. 

Dense, wet, mixed, lowland forest, 40-300 meters; endemic; Iza- 
bal (type from Cerro San Gil, along Rio Frio and tributaries, Steyer- 
mark 416 40; also between Bananera and "La Presa" in Montana 
del Mico). 

Stems solitary, slender, 1-6 meters high, 5-7 mm. in diameter, the internodes 
1.5-3.2 cm. long; leaves 3-5, pinnate, the sheath 6-9 cm. long or rarely more, 
cylindric, briefly oblique at the apex, striate-nerved, the petiole 6.5-12 (rarely to 
33) cm. long, pale below as is the rachis, this 10.5-23 cm. long; pinnae firmly mem- 
branaceous, rich shining green above, paler and dull green below, 3-4 on each side, 
the terminal pair broadly sigmoid, acute to subacuminate, 14-16 (-21) cm. long on 
the upper margin, 5.5-9 cm. on the rachis, one side usually lower than the other, 
with 4-7 primary nerves keeled above, pale and shining below when dry, second- 
ary nerves 2-4 between the primaries, inconspicuous; lower pinnae progressively 
smaller, sigmoid-acuminate, alternate, 11-14 (-21) cm. long, 2.5-3 (-4.5) cm. wide 



236 FIELDIANA: BOTANY, VOLUME 24 

or the lowest only 7.5-9 (-18) cm. long, 1-1.5 (-3) cm. wide, each with a prominent 
central nerve and submarginal nerves and 1-3 pairs of inconspicuous secondary 
ones on each side, at least the central nerve somewhat keeled; inflorescences borne 
below the leaves, arcuate-nodding, subtended by 3 ( cf ) -4 ( 9 ) tubular brown 
spathes, the lowermost inserted about 1 cm. above the base, the uppermost slightly 
exceeding the peduncle in staminate plants, the peduncles 16-21 cm. long, stami- 
nate ones with 5-7 divaricate-pendulous slender green branches (6-) 9-14.5 cm. 
long from a rachis 1-2 cm. long, the strongly ribbed (when dry) flowers 2 mm. long, 
yellowish-green, essentially superficial, rather densely spiralled, the calyx 0.75 mm. 
high, longer than broad, imbricately deeply 3-lobed, the petals shortly united and 
laterally compressed in a linear base, expanded above and connate by their tips, 
the corolla opening by lateral slits, stamens subsessile with anthers entire at the 
apex, sagittate at the base, pistillode 3-angled, truncate; pistillate inflorescence 
with 4-8 slender branches 5.5-13 cm. long from a rachis 1.5-2 cm. long, becoming 
orange in fruit; fruit orange, rather densely spiralled, essentially superficial on the 
branches, sickle-shaped, 1.1-1.4 cm. long, 3-4 mm. in diameter, the perianth nerved 
when dry, calyx 0.7 mm. high, shallowly 3-lobed, petals imbricate, the 2 larger 
2 mm. high, staminodes lacking, abortive carpels adherent to the fruit at maturity; 
seed 9 mm. long, 2 mm. in diameter, the embryo at the middle. 

The fruits of this species are suggestive of the more robust C. neu- 
rochlamys but are distinct in their shape. Staminate and pistillate 
inflorescences of this apparently local species are few-branched, dif- 
fering in this respect from other species allied to it. The foliage is 
distinguished with difficulty from that of C. micrantha and C. Agui- 
lariana. 

Chamaedorea geonomaeformis Wendl. in Otto & Dietr. Allg. 
Gartenz. 20: 1. 1852; Oerst. Ame"r. Centr. pi. 5, f. 1-17. 1863. Nun- 
nezharoa geonomaeformis Hook. f. Bot. Mag. pi. 6088. 1874. Capuca- 
capocha. Figure 41. 

Dense, moist or wet, mixed forest, generally at altitudes of 100 
meters or less or rarely to nearly 600 meters; Izabal; eastern Verapaz 
and Chiquimula; described from plants said to have been grown from 
Guatemalan seed collected by Warscewicz (type material illustrated 
by photograph in Herbarium of Chicago Natural History Museum; 
Negative 29895). British Honduras; Honduras. 

Stems solitary, to 1.7 meters high, 4-7 mm. in diameter, but sometimes flower- 
ing while still nearly stemless; leaves erect-spreading, rich grass green above, grass 
green below, simple, oblanceolate in outline, deeply furcate at the apex in a very 
slender acute angle, the sheath obliquely open above the middle, 5-7 cm. long, the 
petiole 5-15 cm. long, pale-centered below as is the rachis, this 11-22 cm. long; 
leaf blade 6.5-13 cm. long on the upper margin, 6.7-12.5 cm. wide at the tip of the 
rachis, the subscarious margins briefly emarginate or mucronulate at the tips of 
9-12 nerves on each side, toothed between the nerves, the latter pale and promi- 
nent below, not keeled above; spadices solitary from the orifices of the sheaths but 
sometimes persisting below the leaves in fruit, subtended by about 5 closely sheath- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 237 

ing tubular spathes, the uppermost somewhat shorter than to exceeding the erect 
peduncle, the staminate spadices with 1-4 (-6) pendulous green branches, the 
lemon-yellow flowers 2.5 mm. high, sessile in shallow elliptic depressions and rather 
crowded on branches to 18 cm. long but not in distinct groups or rows, the annular 
calyx 0.5 mm. high, broadly 3-lobed, not nerved when dry, the corolla scarcely 
nerved, petals very shortly connate basally, connate at their tips, the corolla open- 
ing by lateral slits, stamens with filaments about equaling the anthers, these entire 
at the apex, briefly separated at the base, the pistillode columnar, truncate; pistil- 
late spadices simple or with 2-3 rachillae 5-13.5 cm. long, erect, becoming some- 
what thickened and orange to vermilion in fruit, the flowers 1-1.2 mm. high, sep- 
arated and sessile in shallow subglobose depressions in a spiral, the calyx of imbri- 
cate, strongly nerved (when dry) sepals, 0.5 mm. high or less, the corolla of 
imbricate nerved (when dry) petals, staminodes lacking, abortive carpels adherent 
to the fruit; fruit blue-black at maturity, globose or nearly so, 7-9 mm. in diameter 
(or to 12 mm. when fresh), falling with perianth attached, the seed to 9 mm. in 
diameter. 

This palm, like some other species, flowers at an early stage of 
development, the first inflorescence sometimes appearing in the axils 
of the third leaf as in C. Ernesti-Augustii, with which it is often con- 
fused. The species is the type of the genus Migandra 0. F. Cook 
(Nat. Hort. Mag. 22: 142. 1943), which was described without Latin 
diagnosis and therefore has no standing. The transfer of Chamae- 
dorea geonomaeformis was suggested but not actually made. 

Chamaedorea Lehmannii Burret, Notizbl. Bot. Gart. Berlin 
11: 857. 1933. Chiquilote (El Progreso) ; Quite (Zacapa). 

Dense, moist or wet, mountain forest, 1400-2600 meters; en- 
demic; Alta Verapaz (type collected in forest near Coban, F. C. 
Lehmann 1374)', El Progreso (Sierra de las Minas); Zacapa (Sierra 
de las Minas). 

Plants small and slender, 2 meters high or less, often flowering when acaules- 
cent or nearly so, the caudex 1 cm. thick or more, densely annulate; sheaths about 
12 cm. long, obliquely open for about half their length, densely imbricate below, 
strongly striate-nerved, attenuate into the base of the petiole, this rather stout, 
15-20 cm. long; leaf blades 50 cm. long or larger, the rachis verrucose on the angles 
and ridges above, the segments regularly disposed, about 8 on each side, narrowly 
lanceolate, very inconspicuously if at all sigmoid, almost equally attenuate to each 
end, the apical ones somewhat broader than the others, the segments with a promi- 
nent keeled central nerve and 2-4 less prominent slender secondary ones, 20 cm. 
long and 2 cm. wide or smaller, the terminal pair sometimes as much as 3.5 cm. 
wide; inflorescences solitary, erect, rising from the orifices of the sheaths, subtended 
by 5 (c?) -6 ( 9 ) tubular slender spathes, the staminate spadix long-pedunculate, 
with about 6 erect branches 18 cm. long or less, the green flowers 3 mm. high, 
essentially superficial, well separated in a loose spiral, the calyx nerveless, rather 
deeply 3-lobed, the petals nerveless, not united at the apex, the anthers not divided 
apically and about as long as the filaments, pistillode columnar; pistillate spadix 



238 FIELDIANA: BOTANY, VOLUME 24 

on a long slender peduncle, the branches 1-3 and 8-11 cm. long, rather stout, 
orange-red in fruit, the flowers rather remote, subglobose, dull green, the calyx 
short-cupular, with broadly rounded lobes, the petals elliptic, obtuse; staminodes 
present, abortive carpels adherent to the corolla in fruit; fruit globose, black, 7 mm. 
long when dry, the seed globose, of about the same diameter. 

This species, as interpreted from plants of the Sierra de las Minas, 
superficially resembles completely pinnate forms of C. adscendens, 
especially in the foliage. It is, however, amply distinct in distribu- 
tion at much higher elevations, in its erect spadices emerging from 
the orifices of the sheaths, in its fewer spathes, and loosely flowered 
staminate spadix with erect branches. 

Chamaedorea micrantha Burret, Notizbl. Bot. Gart. Berlin 
11:749. 1933. 

Dense, wet, mixed, mountain forest, 1200-1600 meters; endemic; 
Alta Verapaz (type collected at unknown locality by Tuerckheim) 
along Rio Frio below Tactic and along Rio Carcha between Coban 
and San Pedro Carcha. 

Plants solitary, erect, the slender caudex about 1 meter high, 7-10 mm. in 
diameter with internodes 1.7-3 cm. long; leaves pinnate, the sheath 10-16 cm. long, 
shortly oblique at the apex, striate-nerved, petiole elongate, 12.5-23 cm. long, pale- 
backed as is the rachis, this 28-38 cm. long with usually 5 pinnae on each side; 
pinnae strongly sigmoid, acuminate, alternate, the largest near the base and there 
15-28 cm. long, 3.5-6.5 cm. wide, the median and submarginal nerves rather promi- 
nent but scarcely keeled above, pale below, the 2-3 secondary nerves on each side 
only moderately conspicuous, the apical pair of pinnae much broader than the 
others (rarely one side divided) with generally 4-5 primary nerves, the secondary 
nerves between them only moderately conspicuous; inflorescences slender, arising 
among or below the leaves, erect but nodding at the tip, the peduncles 24-30 cm. 
long, subtended by 4-5 closely sheathing tubular spathes, the lowest inserted above 
the flattened base of the peduncle, the uppermost often exceeding the peduncle; 
staminate spadices with 12-17 slender spreading-pendulous branches generally 10- 
15 cm. long, but sometimes shorter and sometimes the lower forked, the axis 
3.5-6.5 cm. long, the strongly nerved flowers 3 mm. high, essentially superficial, 
rather densely spiralled, the calyx 1 mm. high, shallowly 3-lobed, longer than 
broad, the petals compressed in a linear base, expanded above and connate at 
their tips, the corolla opening by lateral slits, the stamens with short filaments, 
the anthers bifid at the base, entire at the apex, pistillode 3-angled, subtruncate; 
pistillate spadix with 7-17 spreading or nearly pendulous branches 6-9 cm. long 
from an axis 2.5-5 cm. long, the strongly nerved flowers 2 mm. high, essentially 
superficial, rather loosely spiralled, the calyx 1 mm. high, rather deeply and imbri- 
cately 3-lobed, the petals imbricate, staminodes lacking, pistil ovoid with sessile 
stigmas; fruit not known. 

The shape of the fruit is an important specific criterion in the 
group of species to which material assigned to C. micrantha belongs. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 239 

It is unfortunate that all the specimens studied and agreeing with 
Burret's description are in flower. As it is here interpreted, C. mi- 
crantha seems very close to C. Aguilariana and, in turn, to C. concolor 
from Mexico. Dried foliage has a distinctive appearance pale with 
rather inconspicuous secondary nerves that suggests caution in at- 
tempting to unite the species with another. On the other hand, it is 
not possible to distinguish the species by adequate means and further 
collecting is required to ascertain its status. The leaves, as noted by 
Burret, are more graceful and delicate than those of C. concolor and 
in this respect resemble those of C. falcifera. 

Chamaedorea neurochlamys Burret, Notizbl. Bot. Gart. Ber- 
lin 11: 744. 1933. Docanthe alba 0. F. Cook, Nat. Hort. Mag. 22: 
96. /. 9-11. 1943 (published without Latin description). Pacaya; 
Pacayo; Chilac (Huehuetenango). 

Dense, moist or wet, mixed forest, 700 or mostly 200 meters or 
lower; Alta Verapaz; Huehuetenango; Izabal; Pete"n. Southern 
Mexico to Honduras in Atlantic lowlands. 

Plants solitary, 1.5-4.5 meters high, caudex 1.2-2.5 cm. in diameter; leaves 
few, pinnate, dull dark green above, dull green below, sheaths about 18 cm. long, 
cylindric, obliquely open at the top and there pale-margined (when fresh), striate- 
nerved, the petiole 15-29 cm. long, pale below as is the rachis, this 45-65 cm. long 
with 6-8 pinnae on each side; pinnae regularly spaced and remote except sometimes 
the lower 2-3, narrowly rhombic-sigmoid, acuminate at the apex, narrowed to the 
base, the lower central pinnae longest, 23-33 cm. long, 5-6.5 cm. wide, with promi- 
nent central and submarginal nerves shining and slightly keeled, at least toward 
the base, above and pale and shining below, 4-5 slightly less prominent secondary 
nerves on each side, tertiary nerves fine and numerous, the uppermost pinnae some- 
what broader, 14-16 cm. long on the upper margin, about 5-nerved, the lower 
pinnae smaller; inflorescences arising among or below the leaves, 30-42 cm. long, 
subtended by 5-6 slender tubular appressed spathes, the staminate spadix with 
15-20 or more slender pendulous light green branches 15-20 cm. long from an axis 
4-7 cm. long, the flowers 2 mm. high, yellow, strongly nerved when dry, scarcely 
immersed in the axis, rather densely spiralled, the calyx about 0.5 mm. high, shal- 
lowly 3-lobed, longer than broad, the petals very shortly connate and laterally 
compressed in a linear base, arcuate above to a depressed apex and there connate, 
the corolla opening by lateral slits, the stamens with short filaments, anthers not 
bifid at the apex, pistillode 3-angled, truncate; pistillate spadix erect at anthesis, 
nodding in fruit, simply branched with 10-25 slender ascending branches 9-14 cm. 
long from an axis 6-10.5 cm. long, becoming orange or vermilion in fruit, the flowers 
1.5-2 mm. high, strongly nerved when dry, scarcely immersed in the axis, loosely 
spiralled, the calyx 0.75-1 mm. high, deeply 3-lobed, the lobes rounded, the petals 
imbricate, staminodes lacking, the pistil ovoid with sessile recurved stigmas, the 
abortive carpels adhering to the fruit at maturity; fruit subreniform, obtuse, orange 
(perhaps blackening when dead ripe), about 10 mm. long, 5 mm. in diameter. 



240 FIELDIANA: BOTANY, VOLUME 24 

Chamaedorea neurochlamys, known in British Honduras as "mon- 
key-tail," was described from flowering material collected in the 
vicinity of Cubilgiiitz, Alta Verapaz. A superficial resemblance to 
Chamaedorea oblongata should not be confusing if the nervation of 
pinnae and especially the staminate flowers are observed. 

Chamaedorea nubiuni Standl. & Steyerm. Field Mus. Bot. 
23:202. 1947. 

Moist or wet, mixed, mountain forest, 1500-2900 meters; El Pro- 
greso (type from Sierra de las Minas, hills north of Finca Piamonte, 
toward the summit of Volcdn de Santa Luisa, in cloud forest, Steyer- 
mark 43583a); Huehuetenango (Sierra de los Cuchumatanes, Cerro 
Huitz). Southern Mexico (Chiapas). 

Caudex slender, erect, 1.5-2.5 meters high, 8-10 mm. in diameter, the upper- 
most internodes short, the lower ones as much as 9.5 cm. long; leaves simple, the 
sheath 30 cm. long, tubular, oblique at the apex, laxly appressed, about 13 mm. 
in diameter, striate-nerved, the petiole slender, 9-19 cm. long; leaf blades variable, 
sometimes about 45 cm. long and deeply bifid, the rachis 9.5-15 cm. long, the seg- 
ments 20-33 cm. long on the upper margin, falcately long-acuminate, entire or 
subentire, cuneate-acute at the base, the primary nerves 14-17 on each side, promi- 
nent, keeled above, the secondary ones obscure, or sometimes the blades partially 
to completely pinnate, then with rachis about 33 cm. long, about 13 pinnae on each 
side, the terminal scarcely broader than the others, or the rachis to about 25 cm. 
long, the terminal pair of pinnae broad and about 7-nerved, the lower pairs about 6, 
the pinnae, when developed separately, narrowly falcate, to 21 cm. long, 2 cm. 
wide, the central and a pair of lateral nerves prominent and keeled; peduncles re- 
curved, 25-27 cm. long, 2.5 mm. thick near the apex, the spathes (perfect ones not 
seen) narrowly tubular, striate-nerved, 3-4 mm. thick, acute at the apex, ap- 
pressed; the staminate spadix with about 15 markedly undulate rachillae to 6.5 
cm. long, the laterally compressed flowers subimmersed and rather distant in nar- 
rowly elliptic alveolae in the curves, distributed in a loose spiral but appearing as 
though in opposite lines, the calyx very low, scarcely 0.5 mm. high in bud, mem- 
branous, nerveless, shallowly lobed, the petals nerveless in bud, longer than high, 
the stamens with very short filaments and anthers briefly bifid at the apex, more 
deeply so at the base, pistillode short-columnar, truncate; pistillate spadices in 
fruit about 15 cm. long, the rachis 4 cm. long or sometimes more, the branches 5 
or more, strongly undulate and orange, the flowers subimmersed and rather distant 
in a loose spiral but appearing almost distichous, the alveolae 2.5 mm. long, calyx 
very short, shallowly 3-lobate, the broad lobes subtruncate, nerveless, the petals 
thickened in age, semi-orbicular, nerveless, drying dark brown, staminodes pres- 
ent; fruit at maturity black with a glaucous bloom, ovoid-globose, to about 14 mm. 
long and 11 mm. broad. 

Chamaedorea oblongata Martius, Hist. Nat. Palm. 3: 160. 
1837. Nunnezharoa oblongata Kuntze, Rev. Gen. 2: 729. 1891. 
Chamaedorea lunata Liebm. in Martius, Hist. Nat. Palm. 3: 307. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 241 

1849. Nunnezharoa lunata Kuntze, Rev. Gen. 2: 729. 1891. Cham- 
aedorea fusca Standl. & Steyerm. Field Mus. Bot. 23: 200. 1947. 
Cauquib (Alta Verapaz). Figure 41. 

Moist or wet, mixed, lowland forest, sometimes at least on lime- 
stone, generally 350 meters or lower; Alta Verapaz; Izabal; Pete"n. 
Eastern Mexico to Nicaragua. 

Plants solitary, erect, 1.5-3 meters high, the caudex 1-2.5 cm. thick, the inter- 
nodes unequal, 4-15 cm. long; leaves pinnate, the sheath to 20 cm. long, cylindric, 
oblique at the apex, prominently costate, the petiole 25-31 cm. long, stout, pale- 
backed as is the rachis, this 35-55 cm. long with 6-9 mostly alternate, somewhat 
coriaceous pinnae regularly disposed on each side; pinnae rhombic-lanceolate or 
lanceolate and strongly sigmoid, deep green and glossy above, paler below, the 
middle ones 17-40 cm. long, 3.5-10 cm. wide, narrowly and caudately long-acumi- 
nate or attenuate-acuminate, the primary nerves 3, central and submarginal, these 
not keeled above, yellowish and shining below, the secondary nerves about 3 on 
each side, inconspicuous as are the numerous very fine tertiary nerves, terminal 
pinnae shorter than the succeeding ones and generally narrower; inflorescences 
arising below the leaves, spreading, slender, the peduncles 15-30 cm. long sub- 
tended by 5-7 tubular sheathing striate-nerved spathes; staminate spadix with 9- 
25 or more pendulous slender slightly angled branches to 30 cm. long from an axis 
9-12 cm. long, the superficial or slightly impressed nerveless flowers rather densely 
spiralled, drying black, the calyx shallowly 3-lobed, about 0.5 mm. high, the val- 
vate petals 3-4 mm. high, not connate at the tip but united basally in a short stipe, 
stamens with anthers longer than the filaments and briefly bifid at the apex, pistil- 
lode cylindric, slightly expanded and truncate at the apex; pistillate spadix with 
6-20 or perhaps more stiff branches 9-16 cm. long becoming orange in fruit, the 
nerveless flowers drying black, scarcely immersed in a loose spiral, calyx deeply 
3-lobed, about 1 mm. high, petals imbricate above a very shortly connate base, 
2 mm. high, broadly ovate, acute, drying dark brown in fruit with abortive carpels 
adherent to the smallest petal, staminodes present, pistil with sessile stigmas; fruit 
shining black at maturity, 10-14 mm. long, 7-8 mm. in diameter, ellipsoid or very 
slightly lunate. 

Chamaedorea Pachecoana Standl. & Steyerm. Field Mus. Bot. 
23:202.1947. Pacaya. Figure 41. 

Dense, moist or wet, mixed, mountain forest, 1200-1500 meters, 
or perhaps even higher; endemic (type cultivated in the garden of 
Don Mariano Pacheco Herrarte, Guatemala, said to have come from 
Totonicapan, but perhaps in error, Standley 63104)', Quezaltenango 
(known only from the barranca of the Rio Samala, along the lower 
slopes of the volcanoes of Santa Maria and Zunil). 

Plants very small, 60 cm. high or less, the stems very slender and densely annu- 
late, decumbent and rooting with only the top exposed for about 8 cm., the inter- 
nodes scarcely 6 mm. long; leaves often numerous, small, pinnatisect, mostly 30-60 
cm. long and 8-13 cm. wide, the sheath about 5 cm. long, open to the base, lax, 
striate-nerved, the very slender petioles 7-17 cm. long, somewhat verrucose above, 



242 FIELDIANA: BOTANY, VOLUME 24 

the rachis pale-backed, verrucose on the angles and ridges above; pinnae thin, 
mostly 7-11 on each side, broadly rhombic-lanceolate or oblong-lanceolate, sig- 
moid, 4.5-8 cm. long and 1.5-2 cm. wide or sometimes slightly larger, narrowly 
and falcately long-acuminate, arcuate-cuneate at the base and sometimes very 
minutely auricled with a very narrow basal attachment, the margins shallowly 
toothed toward the tip and often verrucose toward the base, regularly inserted, 
alternate or subopposite, the terminal ones coalescent, sometimes slightly longer 
and broader than the succeeding ones, all the segments 3-nerved, the secondary 
nerves slender and inconspicuous; spadices solitary, erect from the orifices of the 
sheaths or sometimes borne below the leaves, subtended by 5-6 very narrowly 
tubular spathes scarcely 2 mm. in diameter, the upper ones 5-7.5 cm. long, oblique 
at the apex; peduncles very slender, mostly 11-23 cm. long, scarcely more than 
1 mm. thick near the apex; staminate spadix 7-11 cm. long, the rachis very short, 
the branches 1-5, rather densely flowered to the base, the greenish-yellow, subim- 
mersed flowers very numerous, 3 mm. high, the nerveless calyx 2.5 mm. broad, 
shallowly 3-lobate; petals incurved but not connate at the apex, broadly ovate, 
obtuse, not nerved; anthers oblong-ovate, obtuse, pistillode columnar; pistillate 
spadix simple or furcate and 5-10 cm. long, the branches sparsely flowered, rather 
stout, straight, the flowers spirally disposed, subimmersed, the rachis red or orange 
in fruit; calyx 2 mm. broad, shallowly 3-lobate, the nerveless lobes broadly rounded, 
petals pale, rounded-ovate, 2 mm. long, acute, slightly striate-nerved when dry, 
staminodes present, abortive carpels generally adherent to the corolla in fruit; 
fruit black at maturity, globose or ovoid-globose, 6-8 mm. in diameter. 

This is easily the smallest of all the known palms of Guatemala 
and one of the most attractive. Although C. elegans blooms when 
even smaller, it soon attains a much greater size, becoming 2 meters 
high or more. All the plants of C. Pachecoana that we have seen, 
and they are numerous in the forested parts of the barranca of the 
Samala, had only very short emergent caudices, the remainder being 
below ground and often rooting at the nodes. For pot culture, C. ele- 
gans is probably more desirable, since it is more slender and graceful, 
but C. Pachecoana, on the other hand, has usually more numerous 
and somewhat crowded leaves. It is rather remarkable that so un- 
usual a palm should have remained unknown to foreign cultivation, 
especially since it grows almost beside one of the oldest roads of west- 
ern Guatemala, along which various botanists have traveled during 
the past century. 

Chamaedorea parvisecta Burret, Notizbl. Bot. Gart. Berlin 
11:742. 1933. 

Moist or wet, mixed, mountain forest, 1600 meters; Alta Vera- 
paz (type from mountains near Coban, Tuerckheim II. 2185). 

Plants very slender, the caudex arundinaceous, 5-7 mm. in diameter, with 
internodes about 3 cm. long; sheaths elongate, slender, cylindric, oblique at the 
apex; petiole slender, about 30 cm. long; leaf blades about 40 cm. long with 10 or 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 243 

more pinnae on each side of a pale-backed rachis; pinnae small, regularly disposed, 
alternate, lanceolate, not conspicuously sigmoid, long and narrowly acuminate, the 
larger central ones about 15 cm. long, 2 cm. wide (or more?); inflorescences break- 
ing through the sheaths at the nodes(?), the peduncle about 16 cm. long, subtended 
by about 8 spathes, the lowest four of these short and densely congested, staminate 
spadix very slender with about 8 slender branches, the small nerveless flowers semi- 
globose in bud, the calyx low, with 3 broadly rounded, nearly free lobes, petals 
ovate, pistillode columnar; pistillate spadix with about 4 branches 5-5.5 cm. long; 
fruits laxly disposed, seated in distinct depressions in the axis, broadly ovoid, about 
8 mm. long, 7 mm. in diameter; seed with embryo below the middle. 

One specimen, Standley 91426, from mountains along the road 
between Tactic and the divide on the road to Tamahu at an eleva- 
tion of 1500-1600 meters, appears to belong here but the pinnae are 
broader (to 3.5 cm.) than those described by Burret. The latter has 
suggested that C. parvisecta may be no more than a genotypic vari- 
ant of C. pukhra, a possibility borne out by the single collection but 
requiring further study. 

Chamaedorea pulchra Burret, Notizbl. Bot. Gart. Berlin 11: 
741. 1933. 

Moist or wet, mixed, mountain forest, 1550 meters; Alta Vera- 
paz (type collected in Guatemala without special locality, but doubt- 
less from the Coban region; paratype from Coban). 

Caudex slender, about 1 cm. in diameter, the internodes separated; sheath 
elongate, cylindric, thin, becoming dry and persistent, oblique at the apex; petiole 
about 20 cm. long; leaf blade about 45 cm. long, nearly concolorous, papery mem- 
branaceous with 8 or perhaps more pinnae on each side of a pale-backed rachis; 
pinnae lanceolate, slightly sigmoid, regularly disposed with the apical pair broader 
than the others, the larger central pinnae 16-20 cm. long, 2.7-3.5 cm. wide, the 
basal pinnae smaller and closer together, all narrowly acuminate with a conspicu- 
ous midnerve and 2 prominent secondary nerves on each side, tertiary nerves 
slender, numerous, inconspicuous; inflorescences emerging through the sheaths at 
the nodes, the slender peduncles 21-30 cm. long, subtended by about 6 closely 
sheathing spathes; the staminate spadix with about 10 slender branches 10 cm. 
long, the flowers semi-globose in bud and densely spiralled, the nerveless calyx low 
with 3 broadly rounded lobes, petals nerveless, free, ovate, anthers oblong, pistil- 
lode columnar; pistillate spadix with about 10 rather stout branches, these sub- 
angular and ridged when dry, about 6 cm. long, the nerveless flowers slightly im- 
mersed and moderately densely spiralled, the calyx low, shortly cupular at the 
base with 3 broadly rounded nearly free lobes, the ovate petals free, imbricate, 
staminodes present, stigmas sessile. 

No material definitely referable to this species has been studied, 
and the description is taken from Burret. One specimen, tentatively 
referred to C. parvisecta, has the inflorescences breaking through the 
sheaths and the broad pinnae of C. pukhra. It is likely that further 



244 FIELDIANA: BOTANY, VOLUME 24 

collecting will show that these do not represent distinct species, a 
possibility suggested by Burret himself. 

Chamaedorea quetzalteca Standl. & Steyerm. Field Mus. Bot. 
23: 204. 1947. 

Known only from the type, Quezaltenango, dense damp mixed 
mountain forest, along the old road between Finca Pirineos and 
Patzulin, on the lower slopes of Volcan de Santa Maria, 1200-1400 
meters, Standley 87159. 

Plants slender, about 1.5 meters high, the caudex almost 1 cm. in diameter, 
lustrous, the internodes 1.5-3 cm. long; leaf sheaths about 17 cm. long, tubular, 
almost 1.5 cm. thick, finely striate-nerved, the petiole slender, elongate, 2.5 mm. 
thick, the blades pinnate; pinnae about 16 on each side of a pale-backed rachis 
40 cm. long or more, regularly disposed, alternate, thin, deep green on both sur- 
faces, linear-lanceolate, scarcely sigmoid but falcately narrowed to a long-acumi- 
nate tip, the larger middle ones 19 cm. long and 2 cm. wide, gradually narrowed to 
the base, mostly 5-nerved with nerves pale and slightly keeled above, shining and 
pale below when dry, the secondary nerves very slender and obscure, or else 
3-nerved with but a single secondary nerve between each pair of primary ones, 
the uppermost pinnae only 9 cm. long and scarcely 1 cm. wide; peduncles very 
slender, as much as 25 cm. long, 1.5 mm. thick near the apex, the spathes (perfect 
ones not seen) tubular, 3 mm. thick, oblique at the apex; pistillate spadix furcate 
with branches about 8 cm. long, spirally and laxly few-flowered, the rachis very 
slender, somewhat flexuous, the flowers distant, not immersed, the alveolae broadly 
oval, 1.5 mm. long; calyx explanate beneath the fruit, 2.5 mm. broad, shallowly 
3-lobed, the lobes broadly rounded or subtruncate; petals thickened in age, drying 
pale with a dark margin, ovate-rounded or very broadly ovate, sometimes broader 
than long, about 3 mm. long, sometimes 4 mm. wide, not nerved; immature fruit 
subglobose, symmetric, 6 mm. in diameter, broadly rounded at base and apex. 

Chamaedorea Rojasiana Standl. & Steyerm. Field Mus. Bot. 
23:205.1947. Pacaya; Molinillo. Figure 41. 

Moist or wet, mixed, mountain forest, 1200-2600 meters; Quezal- 
tenango (type from Quebrada San Geronimo, Finca Pirineos, lower 
slopes of Volcan de Santa Maria, Steyermark 33479); San Marcos; 
Solola; Suchitepe"quez. Southern Mexico (Chiapas). 

Plants slender, solitary, erect, sometimes acaulescent, with slender prop roots, 
the caudex generally 1-1.5 meters high, 6-8 mm. thick, the internodes 5.5-7 cm. 
or sometimes much shorter; leaf sheaths 7-12 cm. long, 8-14 mm. thick, oblique 
above, prominently striate-nerved, the petiole short, slender, 2.5-5 cm. long or 
rarely somewhat longer but shorter than the sheath, the blades rather small, thin, 
almost concolorous or slightly paler beneath, simple and 7-nerved or usually pin- 
nate with 1-3 pairs of alternate sigmoid pinnae to 13 cm. long and 2.5 cm. wide, 
the lowest smaller than the others, each with a central primary nerve and 2 pairs 
of slightly less conspicuous secondary nerves on each side, the terminal pair with 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 245 

4-6 keeled primary nerves on each side and 1-3, generally 2, secondary nerves 
between them, 13-17 cm. long on the upper margin, 7.5-8.5 cm. along the rachis 
when simple, 4-6 cm. along the rachis when pinnate, then the entire rachis to 12 cm. 
or very rarely to 18.5 cm. long, all the pinnae falcate-acuminate; peduncles 11-20 
cm. long, very slender, scarcely 2 mm. thick near the apex, ascending or spreading; 
spathes 3-4, tubular, appressed, scarcely more than 3 mm. broad, striate-nerved ; 
staminate spadix simple or often composed of 2-3 branches, the branches densely 
flowered to the base, nodding, 7-20 cm. long, 4 mm. thick, the flowers strongly 
nerved when dry, deeply immersed, the calyx 1-1.5 mm. high, connate at the base 
in a short tube, with rounded lobes conspicuously imbricate above, the petals con- 
nate and laterally compressed in a linear base, expanded above and connate at the 
tips, the corolla opening by lateral slits, stamens with short filaments and anthers 
entire at the apex, pistillode columnar, truncate; pistillate spadix simple or com- 
posed of 2 branches, these orange in fruit, 7-9 cm. long, the flowers remote, spirally 
arranged, rather few, deeply immersed, the alveolae oblong or oval, 2.5 mm. long, 
the calyx very short, obscurely 3-lobed; petals irregularly rounded, prominently 
striate-nerved; fruit at first orange, at maturity purple-black, subglobose or ovoid- 
ellipsoid, when dry 7-9 mm. long. 

Related to C. simplex with which simple-leaved forms have been 
confused. The present species is named for Professor Ulises Rojas, 
formerly Director of the Jardin Botanico of Guatemala, oh and near 
whose finca, Pirineos, many of the known specimens of the species 
have been collected. 

Chamaedorea Schippii Burret, Notizbl. Bot. Gart. Berlin 11: 
1038. 1934. 

Moist or wet, mixed forest, often on limestone, 700 meters or 
lower; Alta Verapaz. British Honduras. 

Plants forming colonies by rhizomes, the stems separated, not cespitose, to 
3 meters high or more, to 2.5 cm. in diameter; leaves to 1.2 meters long, regularly 
pinnate, the sheath 18-19 cm. long, tubular for about three-fourths its length below 
an oblique, dark-margined orifice, the petiole 15-25 cm. long, green below as is the 
rachis, the latter at first cinereous furfuraceous becoming glabrate or glabrous, 
8-9 dm. long, with about 30 narrowly linear acute to acuminate straight pinnae on 
each side, all except the somewhat wider terminal 2-nerved pair with a single prom- 
inent central nerve scarcely keeled above, the pinnae when dry slightly plicate; 
pistillate peduncle erect from below the leaves in fruit, slender, 15-20 cm. long, 
subtended by 5 tubular spathes, the upper ones slightly inflated and oblique at the 
apex; the pistillate spadix rather small with about 12 ascending branches, these 
stout, curved-ascending, commonly 6-8 cm. long, red in fruit; fruits globose, black 
at maturity, 8 mm. in diameter, superficial and rather densely disposed, the calyx 
3-lobed, the lobes broadly rounded, petals about 2 mm. long, imbricate, broadly 
ovate, staminodes not apparent, abortive carpels adherent to the fruit. 

The species was described from fruiting material which was com- 
pared with C. graminifolia. The latter, described from cultivated 
material possibly of Guatemalan origin, is very similar but differs, 



246 FIELDIANA: BOTANY, VOLUME 24 

according to Burret, in the elongate fruit with embryo near the mid- 
dle of the seed. 

Chamaedorea simplex Burret, Notizbl. Bot. Gart. Berlin 11: 
758. 1933. 

Wet, mixed forest, often on rocky hillsides, 1350-1440 meters, 
endemic; Alta Verapaz (type collected by Tuerckheim in the Coban 
region; known also from Pansamala, Purulha, and rather frequent in 
the forest between Coban and San Pedro Carcha). 

Plants slender, the stems decumbent and rooting at the base (colonial?) but 
erect for 1-1.5 meters, 5-6 mm. in diameter, green, the leaf scars 2.5-7 cm. apart; 
leaves few, scattered along the upper part of the stem usually for half the length 
of the caudex or less, the sheaths 7-9 cm. long, cylindric, obliquely open for 1.5- 
2 cm. at the apex, green, conspicuously nerved, closely appressed, the petiole 
2.5-14 cm. long, generally shorter than the sheath, leaf blades simple, cleft for 
three-fourths their length or more, the rachis 2-5 cm. long, the lobes broad, diver- 
gent, somewhat sigmoid, short-acuminate, curved along the outer margin, as much 
as 20 cm. long on the upper margin and 5 cm. wide, usually smaller, 6-7-nerved, 
the nerves prominently keeled above; pistillate inflorescences borne in the axils of 
live leaves, with several narrow spathes, 20-26 cm. long above the sheaths, simple, 
the spike 8-9 cm. long, becoming orange in fruit; fruits rather laxly disposed in 
distinct but not deep depressions in the axis, the perianth in fruit prominently 
nerved when dry, with distinctly 3-lobed calyx about 1.5 mm. high, and imbricate, 
somewhat gibbous petals 2-2.5 mm. high, staminodes apparently lacking, abortive 
carpels adherent to the fruit; fruit ellipsoid or oblong, about 9 mm. long and 7.5 
mm. broad, becoming black and shining at maturity, the seed subglobose, 5 mm. 
in diameter. 

All the material assigned to this species is either sterile or in fruit. 
Staminate flowers are not certainly known but are probably very 
similar to those of closely related C. Rojasiana. This is one of the 
neatest and, in some respects, the handsomest of the Guatemalan 
Chamaedorea species, of very distinctive appearance and easily recog- 
nized even in the sterile state. The leaves are small and, according 
to Burret, more deeply divided than in any other species, short and 
with widely spreading divisions. 

Chamaedorea Skutchii Standl. & Steyerm. Field Mus. Bot. 
23: 206. 1947. 

Known only from the type, Quezaltenango, Volcan de Zunil, 
2400 meters, A. F. Skutch 935. 

Caudex slender, 2.5 meters high, 8-12 mm. thick; the internodes 2.5-4 cm. 
long; petiole 22 cm. long or more, the blade pinnate, pinnae 9-11 on each side of 
a pale-backed rachis 22-27 cm. long, the upper pair 4-6-nerved, about 20 cm. long 
on the upper margin to 6 cm. wide, falcate-acuminate, the outer side inconspicu- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 247 

ously crenate-serrate, the other pinnae regularly disposed, mostly alternate, 15- 
21 cm. long, 13-16 mm. wide, not sigmoid, subfalcately long-attenuate, 3-nerved, 
the primary nerves slightly keeled and pale above, pale and shining below, the 
secondary nerves very slender; pistillate inflorescences arising below the leaves, 
the slender peduncle 25-31.5 cm. long, almost 4 mm. thick at the apex in fruit; 
spadix orange in fruit, with about 7 rather markedly undulate branches, especially 
so toward the tip, the nerveless flowers borne in a loose spiral, somewhat longer 
than broad, distinctly sunken in slightly curved elliptic pits bordered on the lower 
side by a liplike margin, the sepals essentially free, imbricate and slightly gibbous 
at the base, about 1 mm. high, petals imbricate, subacute to rounded, about 1.5 
mm. high, becoming brown and narrowly dark-margined in fruit, the one covering 
the abortive carpels scarcely exceeding the calyx, the other two about twice as long 
as the sepals, staminodes absent, abortive carpels adherent to the perianth in fruit; 
fruit obovoid-globose, about 10 mm. high, 8 mm. in diameter; seed globose-ovoid, 
8 mm. long. 

Chamaedorea Skutchii resembles C. nubium in the undulate 
branches of the inflorescence but is distinguished from that species by 
its imbricate pistillate sepals, shallower pits, and lack of staminodes. 

Chamaedorea stenocarpa Standl. & Steyerm. Field Mus. Bot. 
23: 206. 1947. 

Known only from the type; Izabal, Cerro San Gil, 650-900 me- 
ters, Steyermark 41898. 

Plants small, acaulescent, about 60 cm. high; leaves 5 or more, small, 60 cm. 
long or shorter, the open sheaths 4-4.5 cm. long, narrowed upward, striate-nerved, 
the slender petioles 12-20 cm. long, very sparsely verrucose or smooth, the rachis 
14-27 cm. long, pale-backed and verrucose-margined toward the apex below, 
strongly verrucose and angled above, narrowly margined with decurrent bases of 
pinnae, the blades pinnatisect, 14-32 cm. long, 10-20 cm. wide; pinnae 10-14 on 
each side, thin, concolorous, regularly distributed, the middle ones 5.5-12 cm. long, 
1.2-2.5 cm. wide, narrowly oblong-lanceolate, falcately acuminate, narrowed to the 
base, the basal line of attachment about 8 mm. long, the segments 3-nerved, the 
primary nerves slender but prominently keeled, verrucose at least near the base, 
the secondary nerves solitary between the primary ones, very slender and incon- 
spicuous, the lower margin shallowly toothed toward the apex; terminal pinnae 
5-7.5 cm. long, 8-15 mm. wide, 2-3-nerved; pistillate peduncle slender, erect, 
straight, 18 cm. long, 1.5 mm. thick near the apex; spathes 4 or perhaps 5, narrowly 
tubular, almost 3 mm. in diameter, closely appressed, oblique at the apex, con- 
spicuously nerved; spadix rather stout, straight, 4 cm. long, deep orange, the flowers 
rather densely disposed, scarcely immersed, spirally arranged, the alveolae elliptic, 
1.8 mm. long; calyx minute, not nerved, very shallowly 3-lobed; petals not nerved, 
almost 3 mm. long in fruit, rounded-ovate, obtuse, free and imbricate; immature 
fruit oblong-ellipsoid, when dry 8 mm. long and 4 mm. broad, obtuse or rounded 
at the apex, obtuse at the base, abortive carpels adherent to the perianth. 

One of the smallest of Guatemalan palms, noteworthy for the 
numerous, very thin, deep green pinnae, and more especially for the 
very short simple spadix bearing numerous oblong-ellipsoid fruits. 



248 FIELDIANA: BOTANY, VOLUME 24 

Chamaedorea stricta Standl. & Steyerm. Field Mus. Bot. 23: 
207. 1947. 

Moist or wet, dense, mixed mountain forest 1400-1700 meters; 
endemic; San Marcos (Volcan de Tajumulco; type collected on nar- 
row dry ridge above Finca El Porvenir, up Cerro de Mono, Steyer- 
mark 37381; also between Todos Santos Chiquitos and Loma de la 
Paloma). 

Plants acaulescent; leaves 2-3, erect, rather stiff, concolorous or somewhat 
paler beneath, simple, the sheath 8-27 cm. long, open, attenuate upward, striate- 
nerved, rather lax, the petiole 28-35 cm. long, as much as 7 cm. thick; leaf blades 
20-60 cm. long, 13-23 cm. wide, cuneately narrowed at the base, deeply bifid at 
the apex, the rachis 14-30 cm. long, the lobes long-acuminate, directed forward, 
the primary nerves 12-24 on each side, prominent below, the secondary nerves 
numerous, very slender and inconspicuous; peduncles arising from the base of the 
plant, erect, slender, but rather stiff, 80-100 cm. long, 2 mm. thick near the apex; 
spathes 7-10, narrowly tubular, closely appressed, pale green, with very slender 
nerves, the lowest spathes only 7 cm. long, the middle ones 11-28 cm. long, 4 mm. 
in diameter, oblique at the apex; staminate spadix about 24 cm. long, the rachis 
3.5 cm. long, the branches 7, densely flowered almost to the base, slender and some- 
what flexuous, the flowers rather deeply set in narrowly elliptic alveolae 2.5 mm. 
long, the calyx very short, 2.5 mm. broad, very shallowly 3-lobed; petals not 
nerved, free, broadly ovate, obtuse, almost 4 mm. high, erect; branches of the 
pistillate spadix 3-4, erect, rather stout, 8-14.5 cm. long, the flowers remote, spi- 
rally inserted, the rachis 1.5-3 cm. long, red-orange, the flowers scarcely immersed, 
the alveolae elliptic or oval, 2.5 mm. long; calyx 3.5 mm. long, deeply 3-lobed, 
petals not nerved, almost orbicular, subapiculate, 3 mm. long and wide, staminodes 
present and prominent in the fruiting corolla; fruit subglobose, 8 mm. long and 
almost equally broad, broadly rounded; seed globose, whitish brown, almost as 
large as the dry fruit. 

Among the few species with simple leaves, this is outstanding 
because of the greatly elongate, stiffly erect, radical inflorescences. 
The foliage and the closely sheathed elongate peduncles are strik- 
ingly similar to those of C. Ernesti-Augustii. The flowers of both 
sexes, however, are quite different and the pistillate spadix is few- 
branched rather than spicate. 

Chamaedorea tenerrima Burret, Notizbl. Bot. Gart. Berlin 
11 : 858. 1933. Lobia erosa 0. F. Cook, Nat. Hort. Mag. 22: 148. 1943 ; 
26: 20, /. 4. 1947, without Latin description. 

Dense, wet, mixed, mountain forest, 900-1600 meters; Alta Vera- 
paz (type collected above Tactic, on Rio Coban, F. C. Lehmann 1312; 
rather common in this general region, but found elsewhere only at 
Chihob. 

Plants small, generally a meter high or lower, the caudex very slender, 3-7 mm. 
in diameter, green, remotely annulate, often decumbent, the upper internodes 2 cm. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 249 

long; sheath long and cylindric, the strongly rugose petiole 4-8 cm. long, pale below 
as is the rugose rachis, this 6-15 cm. long; leaf blades thin, green on both surfaces, 
broadly obovate in outline and simple with 9-10 pairs of primary nerves, rounded 
to subacute at the base, cleft at the apex with upper margins to 14 cm. long, or 
pinnate with a broad 3-8-nerved apical pair of pinnae and 2-6 pairs of short, 
strongly sigmoid, mostly 3-nerved pinnae with the lower margin produced in a 
prominent auricle, outer margins minutely scarious-roughened, mucronulate and 
emarginate at the tips of the nerves and rather sharply toothed between them; 
staminate spadices solitary at nodes below the leaves, to 12 cm. long, subtended 
by 5 cylindric spathes, with about 10 small slender branches from a rachis 4.5 cm. 
long, the flowers (known only in immature state) superficial in a loose spiral, de- 
pressed-globose, the calyx shortly cupular, 3-lobed, anthers entire at the apex, 
pistillode short, cylindric; pistillate spadix from below the leaves (in fruit) with a 
very slender peduncle 8-15 cm. long, the rachillae 2-3, very slender, 3-6 cm. long, 
recurved and red in fruit; fruits few, seated on superficial widely spaced rounded 
cushions, globose, black at maturity, 6-7 mm. in diameter, the corolla nerveless, 
red-brown when dry without a brown margin, staminodes present, abortive carpels 
apparently adherent to the perianth; seed globose with embryo below the middle. 

The leaves of C. tenerrima are variable. A pistillate plant in 
which the blade is nearly completely pinnate was illustrated by O. F. 
Cook (Nat. Hort. Mag. 26: 19. 1947) under the invalid name Lobia. 

Chamaedorea Tepejilote Liebm. in Martius, Hist. Nat. Palm. 
3: 308. 1849; Bot. Mag. pi. 6030. 1873. Stephanostachys Tepejilote 
Oerst. Vid. Medd. Kjoebenhavn 1858: 27. 1859. Nunnezharoa Tepe- 
jilote Kuntze, Rev. Gen. 2: 731. 1891. Stephanostachys Wendlandiana 
Oerst. Vid. Medd. Kjoebenhavn 1858: 28. 1859. Chamaedorea Wend- 
landiana Hemsl. Biol. Centr. Amer., Bot. 3: 407. 1885. Nunnezharoa 
Wendlandiana Kuntze, Rev. Gen. 2: 730. 1891. Chamaedorea exo- 
rrhiza Wendl. ex Guillaumin, Bull. Mus. Hist. Nat. Paris 28: 542. 
1922. Chamaedorea anomospadix Burret, Notizbl. Bot. Gart. Berlin 
11: 763. 1933. Edanthe veraepacis 0. F. Cook & C. B. Doyle, Nat. 
Hort. Mag. 18: 174, /. 1-9. 1939 without Latin diagnosis (based on 
material from Coban, Alta Verapaz). Pacaya; Ixqui-quib (staminate 
plant), Telom-quib (pistillate plant; Quecchi, fide Cook); Chimp (San 
Marcos; Bojdn (Quezaltenango, San Marcos); Aula-te (Alta Vera- 
paz); Ternero (Coban); Pacaya Grande; Chem-chem (Alta Verapaz). 
Figure 41. 

Moist or wet, mixed forest, 1600 meters or lower, and often 
planted for food or for ornament; Alta Verapaz; Izabal; Suchite- 
pe'quez; Quezaltenango; San Marcos; Pete"n; Huehuetenango; doubt- 
less planted in several other departments Guatemala, Escuintla, 
Santa Rosa, Sacatepquez. Eastern Mexico to Colombia. 

Plants solitary or sometimes clustered but perhaps the individuals distinct, 
erect or sometimes decumbent at the base, often with more or less prominent prop 



250 FIELDIANA: BOTANY, VOLUME 24 

roots, the caudex 2-7 meters high or more, 1.8-10 cm. in diameter, with internodes 
4.5-6.5 cm. long or more; leaves pinnate, large, the sheath cylindric, striate-nerved, 
elongate, oblique at the apex, the petiole stout, elongate, 10-47 cm. long, pale- 
backed as is the rachis, this 6-14 dm. long with 12-25 subopposite pinnae on each 
side at regular intervals; pinnae broadly linear-lanceolate to long-lanceolate, nar- 
rowed and sigmoidly curved at the base, falcately narrowed at the apex, the larger 
median pinnae 16-70 cm. long, 3.5-7.5 cm. wide, thin, the primary nerves generally 
5 but sometimes to 8, keeled above, yellowish and shining below with 4-5 or even 
more secondary nerves interspersed, some or all of these sometimes nearly as prom- 
inent as the primary nerves; especially in large pinnae, the tertiary nerves fine and 
numerous; inflorescences borne below the leaves, solitary at the nodes, subtended 
by 4-5 spathes, the lower short and truncate, the upper rather prominently inflated 
and hoodlike, rather soon marcescent; staminate peduncles 6-17 cm. long with 
18-50 (or more) slender, usually spreading or pendulous branches 7-15 cm. long 
from an axis 6-22 cm. long, the flowers bright yellow, not nerved, somewhat fleshy, 
seated in shallow elliptic depressions, very densely crowded in lines, these number- 
ing 6-7 near the base and often only 4 near the apex or 4 nearly throughout, the 
calyx very short, membranous, annular, about 0.3 mm. high, partially adherent 
to the sides of the alveolae and similarly shaped, petals about 2.5 mm. long, appear- 
ing as though united basally because of crowding but essentially distinct with del- 
toid upper halves inflexed in bud, the tips spreading at anthesis, stamens with 
elongate filaments and small ellipsoid anthers separated at the base, entire at the 
apex, pistillode slender, about as long as the stamens; pistillate peduncles 10-27 cm. 
long with 4-17 or more thickish, straight or flexuous, somewhat angled branches 
3-30 cm. long from an axis 1.5-28 cm. long, the whole becoming orange in fruit, 
the flowers not nerved, slightly immersed in rounded shallow depressions and rather 
densely to loosely spiralled, about 2.5 mm. high, the calyx very small, 0.5-1 mm. 
high, annular, becoming undulate in fruit, the petals imbricate and broadly ovate, 
not gibbous at the base, usually undulate and brown-margined in fruit; staminodes 
lacking, the pistil with sessile stigmas, abortive carpels generally adherent to the 
fruit; fruit ovoid to usually ellipsoid, blue-green maturing black, 13-15 (to 20 when 
fresh) mm. long, 7-8 mm. in diameter. 

Chamaedorea Tepejilote is a most variable species throughout its 
wide range. Segregates have been proposed based principally on size 
and number of parts, but it is difficult to find constant supporting 
characters. Inflorescences may differ greatly in size on a single plant 
in the wild state and tend to be especially large on cultivated plants. 
Size and nervature of pinnae are also variable. Plants from Izabal 
and Alta Verapaz which agree well with the description of C. anomo- 
spadix apparently sometimes form clumps of several stems. Whether 
this is due to the development of rhizomes or to growth of several 
individuals in close proximity is not known. In other respects, dif- 
ferences are of size, not of character. 

In Guatemala, this species is not confined to the Coban region, 
although it is perhaps more abundant there than elsewhere, and cer- 
tainly is cultivated and exploited there for market much more than 
in any other part of Central America. It is the best of the pacayas 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 251 

for eating, but it is by no means the only one used for food. The re- 
gion in which most attention is given to cultivation and sale of this 
particular pacaya is Tactic, where there are large plantations of the 
palm. The cultivated plants commonly are considerably larger and 
bear larger inflorescences than those growing wild in the forests, and 
because the inflorescences are larger, they are more desirable for 
food. 

The best pacayas produced in Alta Verapaz bear a striking simi- 
larity to roasting ears (of maize) covered with their husks, and in size 
they are more like field corn than sweet corn. On wild plants, the 
corresponding organs are often not more than half as large. The 
specific name of this palm is of Nahuatl origin, signifying mountain 
maize. In Guatemala, this name is replaced by the term "pacaya," 
the name given to all Guatemalan pacayas, edible or otherwise. The 
origin of the word "pacaya" is unknown (the Indians of Coban have 
abbreviated it, as they have done with many other words, and call 
the palm "pacai"), but since it is of wide use in Central America as 
well as in South America, one may guess that it is of Quechua deri- 
vation, like the word "chonta," in wide use for various kinds of 
palms, although not so used in Guatemala. 

Chamaedorea Tuerckheimii (Dammer) Burret, Notizbl. Bot. 
Gart. Berlin 11: 766. 1933. Malortiea Tuerckheimii Dammer, op. 
cit. 4: 157. 1904. Kinetostigma Tuerckheimii Burret, op. cit. 11: 
317. 1932. 

Endemic; type collected in the mountains near Coban, Alta 
Verapaz, at 1500 meters, Tuerckheim 8603; sometimes cultivated for 
ornament in the gardens of Coban and introduced into cultivation 
in Europe. 

Plants rhizomatous and probably colonial, dwarf and very slender, sometimes 
a meter high but flowering when not more than 30 cm. high, the green caudex 3-4 
mm. thick, conspicuously annulate, the leaf scars 1-2 cm. apart; leaves few or 
numerous, very small, the sheaths cylindric, oblique at the apex, 5 cm. long, rather 
closely appressed, the petioles 5 cm. long or shorter, the blades cuneate-obovate, 
12-18 cm. long, 3.5-7 cm. wide, rounded or obtuse at the apex, rather coarsely 
crenate, very closely plicate, with about 19 nerves on each side of a pale-backed 
rachis which is verrucose above, deep dull velvety green on the upper surface, 
slightly paler beneath; peduncles about 10 cm. long or more, from the leaf axils, 
the spathes thin, very slender, the uppermost obliquely split on one side, the stami- 
nate spadix small with about 7 slender branches to 4 cm. long from a very short 
rachis, the flowers in bud nerveless, superficial in a loose spiral; pistillate spadix 
simple, thickened and orange in fruit, the fruits superficial in a loose spiral, remote, 
purple-black, ovoid, 8 mm. long or larger. 



252 FIELD IANA: BOTANY, VOLUME 24 

This is one of the smallest palms known and one of the most dis- 
tinct. It is exceptionally attractive because of its dwarf habit and 
the handsome leaves of unique form. In the living state, the leaves 
have a decidedly velvety appearance; they are smaller than those of 
any other Central American palm and probably are the smallest 
leaves produced by any palm in the world. The plant thrives in the 
gardens of Coban, although it seems to be little esteemed. We have 
not seen the plant wild but it is brought into Coban in small quanti- 
ties by the men who peddle orchids. They must know where it grows 
in at least small numbers, but are unwilling to reveal the secret. 
They say that it comes from the "tierra caliente," but if the eleva- 
tion indicated by Tuerckheim is correct, their statements are untrue. 

Chamaedorea vulgata Standl. & Steyerm. Field Mus. Bot. 23: 
208. 1947. Cum (San Marcos) ; Pacaya. 

Moist or wet, mixed, mountain forest, 1300-2350 meters; en- 
demic; Quezaltenango (western slopes of Volcan Zunil); San Marcos 
(type from Volcan de Tajumulco, between Finca El Porvenir and 
Loma Corona, 9 miles northwest of El Porvenir, Steyermark 37735} . 

Plants erect with caudex commonly 2-3 meters high, about 1.5 cm. in diam- 
eter, green, conspicuously annulate, the internodes more than 10 cm. long; leaves 
large, pinnate, the petiole 50-75 cm. long, 7-9 mm. thick, straight; pinnae about 
10 on each side of a green-backed rachis, regularly disposed, mostly alternate, thin, 
deep green, concolorous, linear-lanceolate, the median about 50 cm. long, 7.5 cm. 
wide, not sigmoid, attenuately long-acuminate with nearly straight upper margin 
and falcately curved lower margin, gradually narrowed to the base, 3-3.5 cm. wide 
on larger pinnae, primary nerves 5, keeled above, yellowish, shining and prominent 
below, the secondary nerves 1-3 between the primary nerves, shining below, not 
keeled above, tertiary nerves fine and numerous, the pinnae only slightly decrescent 
in size toward the apex and base of the blade; inflorescences arising below the 
leaves, the peduncles stout, 25-60 cm. long, 2-4 mm. thick at the apex in fruit, 
spreading, subtended by 5 (d") -6(9) tubular subappressed spathes 6-12 mm. 
broad, the lowest only 2.5 cm. long, the uppermost about 21 cm. long, acute, 
oblique at the apex, striate-nerved; rachis of the staminate spadix 4-8 cm. long 
with 9-12 pendulous branches 15-25 cm. long, the green nerveless flowers about 
3 mm. high, superficial, moderately densely spiralled, calyx about 1 mm. high, 
3-lobed and -angled, petals valvate, not connate apically, stamens with rather short 
filaments and elongate anthers generally briefly bifid at the apex, pistillode colum- 
nar, somewhat thickened, 3-angled and truncate at the apex; rachis of pistillate 
spadix 1-4 cm. long with 3-6 branches 15-27 cm. long, becoming orange, thick- 
ened, and pendulous in fruit, the flowers superficial, nerveless, rather densely spi- 
ralled, the calyx about 1 mm. high, 3-lobed, petals rather strongly imbricate, acute, 
staminodes lacking, pistil ovoid with sessile stigmas, the abortive carpels adherent 
to the fruit; fruit probably black at maturity, dull rich green when immature, 
obovoid, 8-9 mm. long; seed globose. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 253 

Because of the strongly nerved large pinnae, this species may be 
confused with C. Tepejilote from which it differs strikingly in char- 
acters of the inflorescence, flowers and fruit. 

The distribution indicated here is rather more limited than that 
in the original description due to inclusion of extraneous elements in 
the latter. 

DOUBTFUL SPECIES 

Chamaedorea Deckeriana (Klotzsch) Hemsl. Biol. Centr. Amer., 
Bot. 3: 404. 1885. Stachyophorbe Deckeriana Klotzsch in Otto & 
Dietr. Allg. Gartenz. 20: 364. 1852. Dasystachys Deckeriana Oerst. 
Vid. Medd. Kjoebenhavn 1858: 25. 1859; Ame>. Centr. 14. pi 6. 
1863. Nunnezharoa Deckeriana Kuntze, Rev. Gen. 2: 731. 1891. 

Described from cultivated plants said to have been sent from 
Guatemala by Warscewicz. Represented in the Herbarium of Chi- 
cago Natural History Museum by a photograph of type material 
(Negative 20757). 

The species is native in Costa Rica and does not appear to occur 
north of that country. 

Chamaedorea Warscewicziana Wendl. Bonplandia 10: 37. 1862. 
Nunnezharoa Warscewicziana Kuntze, Rev. Gen. 2: 731. 1891. 

Described from cultivated plants believed to have been of Gua- 
temalan origin. Represented in the Herbarium of Chicago Natural 
History Museum by a photograph of the type (Negative 20771). 
The species is reported by Burret from Costa Rica and has not ap- 
peared in collections from Guatemala. 

COCOS L. Coconut 

Tall, unarmed palms, the trunk thick, annulate; leaves terminal, pinnatisect, 
the segments lanceolate, the rachis subtrigonous, convex dorsally, the sheath short, 
open, fibrous; spadices inserted among the leaves, erect at first, recurved in age; 
lower spathe short-cleft at the apex, the upper one fusiform, ligneous; flowers mo- 
noecious in the same spadix, bracteate, the staminate solitary or geminate along the 
upper part of the branchlet, the pistillate inserted below, scattered; staminate 
sepals small, acute, erect, valvate, the petals obliquely oblong, acute, valvate; 
stamens 6, included, the filaments subulate, the anthers linear, erect, affixed by 
the bifid base; pistillate flowers much larger than the staminate, ovoid, the peri- 
anth accrescent after anthesis, the sepals thick-coriaceous, erect, imbricate, the 
petals shorter than the sepals, coriaceous, convolute-imbricate, connivent-valvate 
at the apex; ovary 3-celled, attenuate to a short style, the stigmas subulate, 
finally recurved; ovules subbasal, ascending; fruit large, 1-seeded, the style ter- 



254 FIELDIANA: BOTANY, VOLUME 24 

minal, the pericarp very thick and fibrous, the endocarp osseous, 3-pored near the 
base; seed very large, the surface reticulate by the branches of the raphe, the 
endosperm uniform, hollow, the embryo opposite one of the pores. 

As now interpreted, the genus consists of a single species, of al- 
most universal distribution on tropical seacoasts. Numerous Amer- 
ican palms that in the past have been referred to Cocos are now 
placed in other genera. 

Cocos nucifera L. Sp. PI. 1188. 1753. Cocotero; Coc (Quecchi). 

Common along both coasts of Guatemala and abundantly planted 
at low or even at fairly high elevations. 

The coconut, in some regions of the earth a plant of prime eco- 
nomic importance, is too well known to need detailed description. 
The plant is the basis of such a vast literature and it supplies so 
many economic products that a volume would be needed to treat the 
subject fully. In Guatemala and other parts of Central America, 
however, the coconut, although seen almost everywhere at low eleva- 
tions, is of relatively little economic importance. From the coast of 
British Honduras large quantities of ripe coconuts are shipped to the 
United States, and perhaps some are exported from the north coast 
of Guatemala. In Central America the meat of the nuts is used for 
making several popular varieties of sweetmeats, especially the often 
delicious "conserva de coco," a soft candy prepared with either crude 
or refined sugar. More important, perhaps, are the tender green fruits, 
"cocos tiernos" or "cocos de agua" (called "pipas" in southern Cen- 
tral America), which supply a colorless sweet liquid of delicious 
flavor, which always is cool, no matter how torrid the atmosphere. 
This beverage is particularly welcome in areas in which the usual 
water supplies are suspected. 

Ripe coconuts, which are useless for beverage purposes, are car- 
ried to all parts of Guatemala from the lowlands, and their meat is 
used in making sweetmeats. The ripe fruit is called "coco de carne," 
the green fruit "coco de agua." The fruits on sale at Quezaltenango 
showed remarkable variations in shape much more than are ordi- 
narily observed. It is stated that in some parts of Guatemala there 
are trees that produce abnormally small fruits. When the coconut 
germinates the ordinarily empty cavity is filled with the expanding 
embryo, which is white and spongy with a texture almost like that of 
a mushroom. This is considered a great delicacy by some people, 
especially children, and the nuts sometimes are germinated especially 
for food. 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 255 

The coconut is in some respects the most picturesque and attrac- 
tive tree of Central America, infinitely varied in form, but always 
graceful and beautiful. Its groves fringe most of the Guatemalan 
seashores except those bordered by mangrove swamps, and hand- 
some groups of trees extend far inland. The trees thrive and produce 
fruit far from the influence of salt water, but when planted at high 
elevations, as about Guatemala, they seldom produce fruit. Not- 
withstanding its present great abundance in Central America, where 
it spreads freely without the aid of man, the coconut is believed to 
have been imported from the Pacific islands. The date of introduc- 
tion into America is unknown, and it may well have been long prior 
to the arrival of European people. The buoyant fruits, which read- 
ily germinate on seashores, are easily transported by sea currents to 
remote lands. 

Coconut palms, when available, seem to be the favorite nesting 
places of those almost domestic Central American birds, the zanates, 
which, although they may be dear to the hearts of the people, are a 
nuisance in more ways than one, and sometimes a great pest, espe- 
cially to newly planted or sprouting corn. The coconut trees also 
are filled with wasps' nests, but the zanates and wasps live amicably 
together. Predatory animals, especially rats, often climb the trees 
in search of eggs and young birds, especially at night, whereupon a 
regular bedlam prevails, and in the morning the ground is strewn 
with eggshells and dead nestlings. 

For one who wishes to become familiar with the flower and fruit 
structure of the palms, the coconut is the best plant with which to 
begin. Both its flowers and fruits are large, and the structure of the 
fruit is known to almost everyone. The fruit exhibits clearly and on 
a large scale the structure that characterizes even the smallest fruits 
of other palm genera. 

CRYOSOPHILA Blume 

References: Beccari, Webbia 2: 230. 1907 (as Acanthorhiza) ; 
Bartlett, Carnegie Inst. Wash. Publ. 461: 37. 1935. 

Low or tall palms, the trunk cylindric, annulate, bearing near the base spinose 
adventitious roots; leaves flabellate, deeply cleft into 2-4 principal segments which 
are united only at the base, these divided into several acuminate segments with 
filaments between them; petiole unarmed, with a distinct ligule; rachis none; spa- 
dices borne among the leaves, short, 2-3-branched, the ultimate branchlets elon- 
gate, the flowers perfect; s pathos and bracts elongate, coriaceous, deciduous, the 
bractlets setiform, deciduous; flowers globose, perfect, carnose, closed in anthesis, 
subsessile, solitary; calyx of 3 free sepals; petals about equaling the sepals, sub- 



256 FIELDIANA: BOTANY, VOLUME 24 

orbicular, con volute-imbricate; stamens 6, the filaments united to form a tube and 
free only at the apex, the anthers ovate-oblong, dorsifixed near the base; carpels 3, 
completely free, attenuate to a filiform style, stigmatiferous at the apex but not 
thickened, the single ovule in each cell basal, erect; perianth slightly accrescent in 
fruit; fruits medium-sized, globose-oblong, the pericarp scant, spongious-fibrose, 
the endocarp thin-membranaceous; raphe short and inconspicuous, the endosperm 
undivided (not ruminate). 

Eight species are reported for the genus, ranging from southern 
Mexico to Colombia. From Central America five others have been 
described, in Costa Rica and Panama. 

Cryosophila argentea Bartlett, Carnegie Inst. Wash. Publ. 
461 : 40. pi 2-5. 1935. Acanthorrhiza Collinsii O. F. Cook, Nat. Hort. 
Mag. 20: 50. 1941. Cryosophila bifurcata Lundell, Wrightia 1: 53. 
1945. Escoba; Acuum (Maya). 

Moist or wet, mixed, lowland forest, 900 meters or lower; fre- 
quently growing on limestone; Pete"n; Alta Verapaz; Izabal. Cam- 
peche; British Honduras (type from Cornhouse Creek, Manatee 
River, Belize District, H. H. Bartlett 11288). Figure 42. 

Plants slender, as much as 8 meters tall, the trunk about 5 cm. in diameter, 
covered with pungent, simple or branched, spinose rhizoids; leaf blades flabellate, 
about 44-costate, cleft medially almost to the petiole and about 9-cleft on each 
side, the ultimate segments simple and 1-costate, about 3 cm. wide, or 2-4-costate 
and 8 cm. wide, sparsely pilose or glabrate above, silvery beneath, with parallel, 
appressed, white and ferruginous hairs; petioles 70-120 cm. long, appressed-ferru- 
ginous-furfuraceous beneath, green above, 1 cm. thick at the base, 8 mm. broad 
above; ligule triangular, the sheath sordid-lanose, divided apically into long fibers; 
spadix nutant, 60-70 cm. long, subtended by whitish, tomentose or glabrate, stri- 
ate, concave spathes 14-20 cm. long, the peduncle fleshy, floccose-lanate, the whole 
inflorescence laxly twice-branched; flowers glabrous, 4.5 mm. long, 3.2 mm. broad; 
calyx segments connate for a third their length, acute, scarcely equaling the petals; 
petals imbricate, subinvolute, 3 mm. long and broad, in fruit 5 mm. long, gibbous 
outside at the base, carnose, rounded at the apex, the margins thin and hyaline, 
minutely denticulate; ripe fruit yellowish or whitish, subglobose, 12 mm. in diam- 
eter; seed 8-10 mm. in diameter. 

Called "palma de escoba" in Campeche, and "give-and-take" in 
British Honduras. This is presumably the palm that has been re- 
ported from British Honduras as Cryosophila nana (HBK.) Blume 
ex Standley. 

Judging from the description, C. bifurcata does not differ signifi- 
cantly from C. argentea. Leaf segments with briefly bifid apices, the 
apparent distinguishing factor, are usual in Cryosophila. Lundell did 
not contrast his species with C. argentea, to which it would key out in 
Bartlett's treatment. The type was collected on the Punta Gorda- 




FIG. 42. Cryosophila argentea. a, Fruiting branch with leaf attached to stem 
(X '/) b, Fruit (X 1M>)- c, Portion of trunk showing spines (X l /t). 



257 



258 FIELDIANA: BOTANY, VOLUME 24 

San Antonio Road near Jacinto Creek, Toledo District, British Hon- 
duras, Gentle 1+972. 

DESMONCUS Martius 

References: M. Burret, Die Palmengattung Desmoncus Mart., 
Repert. Sp. Nov. 36: 197-221. 1934; H. H. Bartlett, Certain Des- 
mond (Palmae) of Central America and Mexico, Journ. Wash. Acad. 
Sci. 25: 81-88. 1935. 

Very spiny, scandent palms, with elongate, flexuous, annulate stems, the spines 
straight or hooked; leaves distant, subsessile, pinnatisect, the segments opposite 
and alternate or scattered, acuminate, shortly narrowed at the base, the rachis of 
the leaf ending in a long, whip-like portion armed with numerous abruptly reflexed, 
stout spines, the petiole short, the sheath elongate, produced above as an ocrea; 
spadices solitary, small, the branches slender, fastigiate, flexuous, the flowers mo- 
noecious, on the same spadix, bracteate and bracteolate, greenish; spathes 2, the 
lower one short, coriaceous, open at the apex, the upper subligneous, cylindric, 
cleft ventrally; staminate flowers numerous on the upper part of the branches, the 
lower flowers pistillate, solitary or subtended on each side by staminate flowers; 
staminate calyx minute, membranaceous, trifid or trigonous, the 3 petals thicker, 
obliquely ovate or lanceolate, acute or acuminate, valvate; stamens 6, included, 
the filaments subulate or filiform, the anthers linear, bifid at the base, erect; pistil- 
late flowers much smaller than the staminate, coriaceous, the perianth slightly 
enlarged after anthesis, the calyx cupular, 3-5-denticulate at the apex, the corolla 
urceolate, truncate or 3-dentate at the apex; ovary ovoid, 3-celled, the style short, 
with 3 acute, revolute stigmas; fruit small, red, ovoid, obovoid, or globose, 1-celled 
and 1-seeded, the pericarp thin, fleshy, the endocarp crustaceous or osseous, reticu- 
late-veined, apically or above the middle 3-pored, the pores stellately fibrous; 
endosperm undivided, the embryo opposite one of the pores. 

About 50 species, ranging from southern Mexico to the West 
Indies and South America. A few others besides those listed here 
are found in southern Central America. The plants are easy of rec- 
ognition because of their climbing habit, and the long naked leaf tips, 
armed with abruptly reflexed spines. They are all too abundant in 
the lowlands of northern Guatemala, as along the Atlantic coast of 
Central America generally, often forming dense tangles that reach to 
the tops of tall trees, and conspicuous because of the bunches of 
bright yellow fruits. The stems sometimes reach a length of several 
hundred feet. The plants are a constant menace to a person traveling 
through the forests, because the stems often dangle loosely from the 
supporting trees, and the long whiplike leaf tips with their sharp rigid 
spines inflict painful wounds and are dangerous to the eyes. The 
strong and flexible stems, like rattan, are sometimes used as a substi- 
tute for rope, and around Lake Izabal strips of the bark are used for 
making the celebrated bayal baskets, one of the distinctive products 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 259 

of that region. The name "bayal" presumably is applied to all local 
species, which are much alike except in minor details. The name 
"balaire" is given in Honduras to one or more of the species of Des- 
moncus. The following key to species is taken from that published 
by Bartlett, except for the addition of D. leiorhachis. 

Corolla of pistillate flower containing at one side of the ovary a fimbriate, oblong, 
basally attached scale as long as the corolla lobes; leaflets small, about 1 cm. 

wide D. anomalus 

Corolla of pistillate flower containing no scale like that described above; leaflets 
more than 2.5 cm. wide. 

Rachis of the leaf unarmed D. leiorhachis 

Rachis of the leaf aculeate and aciculate. 

Leaf rachis, even when short-aciculate, also armed below the middle with a 
few distant, retrorse or subretrorse spines which are the longest ones on 
the plant. 

Lower spathe aciculate; pistillate flowers with annuliform-cupulate calyx, 

very obtusely 3-apiculate, and a corolla broader than high, almost 

evenly truncate but sharply though minutely apiculate . . . .D. Schippti 

Lower spathe almost or wholly unarmed; pistillate flowers with an acutely 

triangular-cupulate calyx and corolla higher than broad, the margin 

3-apiculate from deep, rounded sinuses D. quasillarius 

Leaf rachis sparsely to densely aciculate, with spines no longer than those on 
other parts of the plant, and not clearly dimorphic. 

Lower spathe almost completely unarmed D. uaxactunensis 

Lower spathe aciculate D. ferox 

Desmoncus aculeatus Wendl. Ind. Palm. 20. 1854. Atitara 
aculeata Kuntze, Rev. Gen. PI. 727. 1891. 

According to Burret (Repert. Sp. Nov. 36: 221. 1934), this is a 
nomen nudum and not referable to any known species. The name 
was based upon Guatemalan material. 

Desmoncus anomalus Bartlett, Journ. Wash. Acad. Sci. 25: 
84. 1935. 

Type from Secanquim, Alta Verapaz, Cook & Doyle 97. Known 
only from the type material. 

Stems very slender; ocrea of the leaf sheath 12 mm. in diameter, 22 cm. long, 
furfuraceous, appressed-aciculate; petiole 1.5 cm. long, weakly short-aciculate; 
rachis 165 cm. long, furfuraceous, sparsely armed on the upper surface with spines 
as much as 12 mm. long; leaflets about 19 on each side, slightly aciculate at the 
base, the lower ones 22 cm. long and 7 mm. wide, the middle ones 10-12 mm. wide, 
the uppermost caudate-acuminate; middle leaflets terminating in long portions 
bearing about 10 pairs of reflexed spines 2.5-6 cm. long; terminal cirrhus of the 
leaf unarmed; lower spathe 28 cm. long, 14 mm. wide, sparsely and shortly ap- 
pressed-aciculate; peduncle 6 cm. long, armed with spines 4-10 mm. long; rachis 
whitish, 20 cm. long, sparsely ferruginous-furfuraceous, the flower-bearing branches 



260 FIELDIANA: BOTANY, VOLUME 24 

about 35, very flexuous, the lower ones 12 cm. long; lower flowers ternate; calyx 
subannuliform or rounded-triangular, the angles rounded; corolla very obtusely 
apiculate, bearing within on one side a scale about 2.2 mm. long and as long as 
the corolla lobes, the scale fimbriate at the apex. 

Desmoncus ferox Bartlett, Journ. Wash. Acad. Sci. 25: 87. 1935. 
Bayal. 

Wet mixed lowland forest, at or little above sea level; Pete*n (type 
from Tikal, H. H. Bartlett 12584; collected also at La Libertad); 
Izabal; Alta Verapaz. British Honduras. Figure 43. 

Stems slender, 2 cm. thick, covered by the sheaths, often 10 meters long or 
more; ocrea of the sheath 25 cm. long, ferruginous-furfuraceous, densely armed 
with slender, black spines 12-22 mm. long; free portion of the petiole 2-3 cm. long; 
leaf rachis sparsely ferruginous-pubescent, armed on the upper side with spines 
20-28 mm. long or longer, about 2.5 meters long, the cirrhus 1 meter long; leaflets 
about 24 on each side, the lowest ones 30 cm. long and 1.5 cm. wide, attenuate into 
a filiform tail 8 cm. long, the middle ones 4 cm. wide, acuminate, the uppermost 
about 36 cm. long and 4.5 cm. wide, minutely black-pubescent, armed above near 
the base with numerous spines 3 cm. or more long; leaflets bearing hooked terminal 
spines only 2, the pairs of spines 10, rigid, the lowest 9 cm. long; lower spathe 22 
mm. wide, the closed portion more than 21 cm. long, the open portion 13 cm. long, 
ferruginous-furfuraceous above and moderately aciculate, the upper spathe fusi- 
form, 40 cm. long, 3 cm. thick, aciculate, the spines 18 mm. long or less; peduncle 
7 cm. long, unarmed below, aciculate above, the flowering axis 24 cm. long, un- 
armed, ferruginous-pubescent, the branches about 40, the lower ones 15 cm. long; 
pistillate flowers 3 mm. broad, 2.5 mm. high, depressed-globose; calyx subrotund, 
annuliform, the margin shortly 3-apiculate, 3.3 mm. broad; corolla cyathiform; 
staminate calyx membranaceous, 3.5 mm. in diameter, triangular, the angles pro- 
duced and acuminate, the petals deltoid, 8-10 mm. long, acuminate. 

Desmoncus leiorhachis Burret, Repert. Sp. Nov. 36: 203. 1934. 

Type from Rio Grande, British Honduras, Schipp S51 7a. Doubt- 
less extending into Pete"n or the forests of the North Coast of Guate- 
mala, and sterile material from Alta Verapaz probably is referable 
here. 

Leaf sheath about 2.5 cm. in diameter, furfuraceous, armed with black spines 
5 mm. long; leaves 1.25 meters long, the cirrhus 50 cm. long, unarmed or with a 
very few black spines, the hooks about 9 on each side, stout; petiole very short or 
none, the rachis unarmed; leaflets 26 on each side, the lowest somewhat smaller 
and narrower, lanceolate, narrow-acuminate, the largest 20 cm. long and almost 
2.5 cm. wide, unarmed and glabrous; free portion of the peduncle 10 cm. long, 
furfuraceous and black-setulose; upper spathe 20 cm. long, narrowly short-acumi- 
nate at the apex, 6 cm. broad, armed with black spines about 7 mm. long; rachis 
of the inflorescence 8 cm. long, with about 20 branches, the longest 10 cm. long; 
staminate flowers 1 cm. long, the calyx teeth acuminate, the petals lanceolate, 
acuminate; pistillate calyx annular, the corolla urceolate, the teeth short, broad, 
subapiculate. 




S. H. Grove 



FIG. 43. Desmoncus ferox. a, Fruit attached to rachis (X M). b, Inflo- 
rescence with pistillate flowers in position, the staminate fallen ( X *A). c, Spathe 
of inflorescence ( X V*)- d, Middle portion of leaf ( X 2 A). e, Apex of leaf ( X 2 A). 

261 



262 FIELDIANA: BOTANY, VOLUME 24 

Desmoncus quasillarius Bartlett, Journ. Wash. Acad. Sci. 25: 
85. 1935. Bayal. 

Moist or wet, mixed, lowland forest, 400 meters or lower; Pete'n; 
Alta Verapaz; Huehuetenango; sterile material from Izabal also may 
belong here. British Honduras, the type from Corozal District, 
P. E. Gentle 348. 

Stems 3 cm. thick; ocrea of the sheath 15 cm. long, armed with black spines 

1 cm. long or less; petiole about 2 cm. long, sparsely aciculate, the rachis 2 meters 
long, sparsely ferruginous-furfuraceous, armed above with two kinds of spines, a 
few of them compressed and almost 14 mm. long, others shorter, all spreading or 
subascending, armed beneath below the middle with a few stout, reflexed spines 

2 cm. long, produced above into an unarmed cirrhus; leaflets about 22 on each 
side, the lowest 23 cm. long and 14 mm. wide, aciculate above near the base, the 
middle ones 30 cm. long and 3 cm. wide, acuminate, two of the leaflets produced 
into an armed appendage, this bearing 9 pairs of recurved spines 7 cm. long; lower 
spathe unarmed or sparsely aciculate, 14-18 mm. wide, the open portion 7 cm. 
long, the upper spathe fusiform, 27 cm. long, 22 mm. thick, ferruginous-furfura- 
ceous, short-aciculate, the spines 2-4 mm. long, densely spreading-aciculate above; 
peduncle 5 cm. long, spreading-aciculate; axis of the inflorescence unarmed, the 
branches about 35; pistillate flowers conic or subcylindric, the calyx cupuliform, 
2.5 mm. broad, 1 mm. high, the margin minutely triapiculate, the corolla 3.5 mm. 
high, tri cuspidate at the apex; staminate flowers 10-12 mm. long, the calyx acutely 
triapiculate, the angles acute. 

Called "basket tietie" and "basket whist" in British Honduras. 

Desmoncus Schippii Burret, Repert. Sp. Nov. 36: 202. 1934. 
D. Lundellii Bartlett, Journ. Wash. Acad. Sci. 25: 84. 1935 (type 
from El Paso, Pete'n, Lundell 1555 in April, 1932). 

Type from Rio Grande, British Honduras, at 45 meters, Schipp 
S517. Bayal. Known definitely only from these collections. 

Stems scandent to a height of 20-25 meters, 2.5 cm. in diameter; ocrea of the 
sheath 18 cm. long, armed with black spines 16 mm. long or less; free portion of the 
petiole 2 cm. long, armed with two kinds of spines, the shorter ones about 6 mm. 
long, the longer ones 3.5-5 cm. long; rachis 1.75 cm. long; leaflets 20-25 on each 
side, lanceolate, acute, the lowest about 17 cm. long and 2.5 cm. wide, the middle 
ones 32 cm. long and 3.5 cm. wide, aciculate above near the base, the longer spines 
3-4 cm. long; cirrhus sparsely long-aciculate, its larger, reflexed spines 3.5-3.8 cm. 
long; lower spathe almost glabrous, armed at the apex with sparse, appressed 
spines, the upper spathe fusiform, 22-30 cm. long, 3-4 cm. in diameter, densely 
armed with spreading, black spines 12 mm. long; peduncle 4-10 cm. long, acicu- 
late; branched portion of the inflorescence 18-20 cm. long, aciculate only at the 
base, with about 30 simple branches, the longer ones 10 cm. long; staminate calyx 
3.3 mm. broad, the petals deltoid, 9 mm. long, acute; pistillate calyx 3 mm. in 
diameter, annuliform, the corolla cyathiform, 2.5 mm. high; fruit ovoid, 13 mm. 
long, 11 mm. in diameter. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 263 

It is not certain that the two names listed are synonymous. 
There is some question as to the validity of the various species here 
recognized, but this can be determined only when a sufficient amount 
of material is available for study. 

Desmoncus uaxactunensis Bartlett, Journ. Wash. Acad. Sci. 
25:86. 1935. 

Type from Uaxactun, Pete"n, Bartlett 12576. Known only from 
the original collection. 

Stems 5-15 meters long, 4-5 cm. thick (including the sheaths); ocrea of the 
sheath up to 30 cm. long, furfuraceous-pubescent, armed with black spines 4-18 
mm. long; free portion of the petiole 2 cm. long, short-aciculate, the rachis 2.25 
meters long; leaflets about 25 on each side, the lowest 25 cm. long and 2 cm. wide, 
the middle ones 30 cm. long and 4 cm. wide, acuminate, armed above near the base 
with a few spines 2 cm. long or less; two of the middle leaflets produced into whip- 
like appendages bearing 9 pairs of retrorse spines 1-6 cm. long, the cirrhus spine- 
armed only at the apex, the spines about 2 cm. long, or almost suppressed; lower 
spathe semicylindric, 40 cm. long, 2 cm. broad, brown-lepidote, the upper spathe 
fusiform, 28 cm. long, 4 cm. thick, furfuraceous-pubescent and densely armed with 
black spines; free portion of the peduncle 5 cm. long, armed with spines 3-5 mm. 
long, the branched portion unarmed, 20 cm. long, with 40-50 simple branches 
4-12 cm. long; fruit ovoid, 1.5 cm. long, 1.2 cm. in diameter, minutely apiculate. 

ERYTHEA S. Watson 
Reference: H. E. Moore, Gent. Herb. 8: 215. 1951. 

Erect, single-stemmed, medium to tall, hermaphrodite palms, the trunk naked 
and fibrous-roughened below, clothed with a shag of deciduous leaf sheaths above; 
leaves palmate, the sheath soft fibrous, unarmed; petiole with margins denticulate 
to strongly dentate or rarely nearly smooth, a ligule present at the base of the 
blade above, absent below; leaf blades flabellate to nearly orbicular in outline, very 
shortly costapalmate, regularly divided to the middle or beyond into numerous 
1-nerved segments with bifid apices; inflorescences interfoliar, elongate, with sev- 
eral once- to twice-branched primary divisions, the spathes tubular, sheathing the 
peduncle and primary branches; flowers small, in groups of 3-2 (-1) on slightly 
elevated bracteolate cushions, the sepals 3, free and lightly imbricate, petals 3, 
free or shortly united at the base, valvate, foveolate, stamens 6, the filaments 
united in a ring at the base and adherent to the corolla, becoming attenuate toward 
the anthers, carpels 3, glabrous above, tomentose below at least as fruit develops, 
lightly connate below, the styles connate in a conic beak stigmatic at the apex, 
ovules solitary, basal, erect; fruit subglobose, mostly 20 mm. long or more with 
abortive carpels basal, the stigmatic scar subapical, exocarp smooth at maturity, 
lightly tomentose when young, mesocarp fleshy, endocarp thin and often adherent 
to the seed; seed with narrow elongate raphe, the endosperm homogeneous with a 
deep intrusion of the integument below the raphe, embryo lateral. 

The genus is composed of about 8 species and is closely allied to 
Brahea. One species is known from Guatemala with possibly a sec- 



264 FIELDIANA: BOTANY, VOLUME 24 

ond from British Honduras. The latter is not well enough known to 
be transferred to the genus with certainty and is treated as a doubtful 
species under the original name, Paurotis Schippii. 

Erythea salvadorensis (Wendl. ex Beccari) H. E. Moore, Gent. 
Herb. 8: 217. 1951. Brahea salvadorensis Wendl. ex Beccari, Webbia 
2: 105. 1907. Acoelorraphe salvadorensis Bartlett, Carnegie Inst. 
Wash. Publ. 461 : 32. 1935. Acoelorraphe Cookii Bartlett, I.e. Palma. 

Open dry rocky hillsides, sometimes in Juniperus forest, or in oak 
and pine forest, 800-1600 meters; Baja Verapaz (type of A. Cookii 
collected between Santa Rosa and Salama, 0. F. Cook & C. B. Doyle 
275; also between Santa Rosa and Purulha); Zacapa (Sierra de las 
Minas; region of Santa Rosalia). El Salvador (the type from Con- 
chagua ["Contshagna"] on the Golfo de Fonseca) ; Honduras. 

A medium-sized palm, sometimes 6 meters high, often acaulescent, frequently 
forming colonies; leaf sheaths large and thin, glabrous, rufous, breaking up into 
numerous fibers; petiole elongate, to 4 cm. broad at the base, 2 cm. at the middle, 
the margins with many acute, incurved teeth, when young furfuraceous-lanate; 
ligule bilobate, chartaceous, the lobes triangular-attenuate; blades with about 70 
plicate divisions on each side, when young floccose-furfuraceous on the nerves, the 
segments about 85 cm. long and 1-2 cm. wide, the rachis extending into the blade 
for 12-14 cm.; spadix 1-1.5 meters long or more, the spathes more than 7, the low- 
est one more than 35 cm. long, opening obliquely at the apex, rather rigid, red, sub- 
tending a partial inflorescence more than 70 cm. long; ultimate flowering branchlets 
slender, 8-18 cm. long, griseous-tomentose; flowers very shortly pedicellate, the 
pedicels concealed by the calyx, this about 1.5 mm. high, 2 mm. in diameter, 
sparsely pubescent at the middle and base; corolla 3.5 mm. long, 2 mm. broad, the 
segments reflexed, dorsally subtomentose or glabrous throughout; pistil glabrous 
at anthesis but tomentose below and the young fruits densely white-tomentose, 
more than 1 cm. long. 

This is apparently the handsome palm that is so conspicuous on 
the dry open hillsides of Baja Verapaz (not Alta Verapaz as stated 
in the original description of A. Cookii), growing with oaks, pines, 
Juniperus comitana, and other trees. It commonly forms small iso- 
lated colonies, some of which consist only of low plants, 2 meters high 
or less. The leaves of this palm are much used at the picturesque 
village of Conchagua in El Salvador for making the jaunty hats pe- 
culiar to that settlement. 

EUTERPE Gaertner 
Reference: Burret, Bot. Jahrb. 63: 49. 1929. 

Solitary or cespitose, unarmed, monoecious palms with pinnate leaves and in- 
terfoliar or infrafoliar inflorescences subtended by two spathes, the outer short, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 265 

ancipitous, and open at the apex, the inner elongate, cylindric in bud with gener- 
ally a pronounced acute beak; peduncles short or elongate, the branches few to 
mostly numerous, spreading; flowers in groups of a central pistillate and two lateral 
staminates, the staminate flowers with imbricate sepals, valvate petals, 6 stamens 
with elongate filaments straight or geniculate at the apex, and an often trifid pistil- 
lode; pistillate flowers sessile or sunken in the axis, the sepals and petals imbricate, 
staminodes ordinarily present and dentiform, pistil unilocular and uniovulate; fruit 
usually subglobose with lateral or subapical stigmatic scar, the exocarp smooth or 
minutely roughened (in drying), mesocarp thinly pulpy with flat fibers, endocarp 
thin; seed globose with homogeneous or ruminate endosperm, the embryo basal. 

About 60 species are known for the genus, which ranges from 
Guatemala to southern Brazil. In addition to the single species that 
follows, Euterpe oleracea Mart, has been reported vaguely and not at 
all reliably from British Honduras, where it is surely not native. The 
palms are said to be known there as "monkey-tail palm" and "moun- 
tain cabbage palm." 

Euterpe macrospadix Oersted, Vid. Medd. Kjoebenhavn 1858: 
41. 1859. Euterpe leucospadix Wendl. ex Hemsl. Biol. Centr. Amer., 
Bot. 3: 401. 1885, name only. Euterpe panamensis Burret, Notizbl. 
Bot. Gart. Berlin 11: 865. 1933. Plectis Oweniana 0. F. Cook, Bull. 
Torrey Club 31: 352. 1904; Smithsonian Misc. Coll. 98 (7); 22, pi. 4, 
7, 20-26. 1939. Ternera; Halaute (Quecchi). 

Wet or moist, mixed forest, often on limestone, to 1000 meters; 
Alta Verapaz (type of Plectis Oweniana collected from steep slopes 
and summits of limestone mountains northwest of Panzos near Finca 
Sepacuite, between Senahu and Cabajon, 0. F. Cook & R. F. Griggs 
87). British Honduras to Panama. 

Trunks smooth, annulate, 25 meters tall or more, about 50 cm. in diameter 
1 meter above the thickened base and mass of red-brown exposed or prop roots; 
leaf sheaths about 1.5 meters long or more forming a distinct crown-shaft, the 
brown-scaly petiole to about 50 cm. long, 2.7 cm. wide at the apex, the ascending 
and arching brown-scaly rachis 2.67 meters long or even more, with 60-75 drooping 
pinnae on each side, the lower ones widely spaced, the remainder more regular and 
about 4 cm. apart; pinnae linear, the central ones to nearly a meter long, to 4.5 cm. 
wide, all minutely brown-dotted with membranous scales along the fine nerves 
below and with conspicuous red-brown larger scales on the prominent midnerve 
and 2 pairs of rather prominent lateral nerves below; inner spathe densely ap- 
pressed brown lepidote-tomentose, the peduncle short and stout, the rachis to 40 
cm. long with as many as 70 spreading densely white-tomentose branches to 70 cm. 
long, the lower subtended at first by conspicuous deltoid bracts; staminate flowers 
about 4.5 mm. long, the sepals 2.5 mm. long, acute, scarious-margined, petals 
acute, stamens with filaments scarcely geniculate at the apex, pistillode deeply 
trifid; pistillate flowers sunken in the axis, the fruit inserted in prominent pits, 
subglobose with subapical stigmatic scar, about 1 cm. in diameter when mature, 
the seed with homogeneous endosperm. 



266 FIELDIANA: BOTANY, VOLUME 24 

Plectis Oweniana was described as the only species in the genus, 
but the type, when compared with specimens agreeing with Oersted's 
description and collected near the type locality, does not seem to be 
distinct from Euterpe macrospadix. The species is apparently rare 
in Guatemala. 

GEONOMA Willdenow 
Reference: Burret, Bot. Jahrb. 63: 144. 1930. 

Plants slender, low or rather tall, unarmed, the trunk annulate; leaves terminal 
or alternate, the blades entire, bifid at the apex, or pinnatisect (in the Guatemalan 
species), the segments acuminate, the sheath tubular, short or elongate; spadices 
suberect or decurved, simple, furcate, or paniculately branched, slender or stout; 
spathes 2, usually caducous before anthesis, the lower one incomplete, truncate, 
concave, the upper compressed or fusiform; flowers small, glumaceous, unisexual, 
immersed in spirally disposed pits in the branches, in groups of 3-1, the central 
flower pistillate (or abortive) maturing after the lateral staminate flowers; sepals 
of the staminate flower slightly imbricate below, the petals valvate, connate below; 
stamens 6, the filaments connate below into a tube, the anther cells free; pistillate 
sepals narrow, slightly imbricate below, the petals valvate, connate below; ovary 
1-celled, the ovule ascending, the style lateral, long, the 3 stigmas subulate; fruit 
small, the pericarp usually thin, with slender fibers; endosperm uniform. 

One of the largest genera of palms, with about 170 species, widely 
distributed in tropical America. In Central America, at least, the 
plants are confined to deep wet primeval forest, and do not establish 
themselves in secondary forest. Most of them are small, graceful, 
and handsome plants. 

Pits of the spadix branches in which the flowers are inserted more or less bilabiate, 
the upper margin somewhat produced or conspicuously angulate, the pits 
spirally or verticillately arranged; lower lip of the pit incised or entire. 
Spadix branches slender, the spadix 2-3-branched; pits ternate in alternating 

whorls G. longepetiolata 

Spadix branches stout. 

Spathes narrow, long-tubular, the upper one inserted high above the lower, 

thin G. polyneura 

Spathes broad, inserted close together, firm and coriaceous G. Seleri 

Pits gradually ascending and graduated into the surface of the branch, not obvi- 
ously bilabiate. 

Pits 8-10 mm. apart; leaf segments 5-11 on each side G. membranacea 

Pits only 3-6 mm. apart. 

Leaf segments about 25 pairs, few-nerved G. binervia 

Leaf segments usually 3-7 pairs. 

Leaf segments 3 pairs, all of them several-nerved G. mexicana 

Leaf segments 5-7 pairs, partly 1-nerved, partly several-nerved. 

G. leptoclada 

Geonoma binervia Oerst. Vid. Medd. Kjoebenhavn 1858: 33. 
1859. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 267 

Moist or wet, dense, mixed lowland forest, at or little above 
(about 150 meters) sea level; Pete"n; Alta Verapaz; Izabal. Southern 
Mexico; British Honduras, along the Atlantic coast to Panama. 

Plants 3-6 meters high, the trunk 5-7 cm. thick, conspicuously annulate; leaves 
ascending, unequally pinnatisect, the petiole 30-60 cm. long; the blades as much as 
2 meters long; leaf segments about 25 pairs, variable in width, glabrous or some- 
times sparsely scurfy on the nerves, 50-60 cm. long, the narrowest ones about 1 cm. 
wide, the larger ones 7-8 cm. wide or the terminal ones 12 cm. wide or more, nar- 
rowly long-attenuate at the apex, obliquely attached and not narrowed below; 
spadix much branched, the long peduncle subtended at the base by 2 conspicuous 
erect valves 15-20 cm. long; inflorescences 60-75 cm. long, pendent, diffusely 
branched, sometimes erect, the ultimate branchlets somewhat pubescent, dark red 
or maroon, 20-30 cm. long, 2-3 mm. thick, the flowers minute, purplish; staminate 
flowers about 4 mm. long, usually somewhat exserted at anthesis; pistillate flowers 
smaller, the 3 stigmas exserted; fruit globose-oblong or globose, 4-6 mm. long, 
glabrous, brownish black at maturity, or sometimes black and lustrous. 

Known in Honduras by the names "manaca" and "pacuca." 

Geonoma leptoclada Burret, Notizbl. Bot. Gart. Berlin 11: 
863. 1933. Cum (Quezaltenango). 

Moist or wet, mixed, mountain forest, 1000-2000 meters; so far 
as known, endemic, but to be expected in Chiapas; Solola (Volcan 
de Atitldn); Quezaltenango (type from Volcan de Santa Maria, at 
2000 meters, F. C. Lehmann 1613; collected several times on the 
lower slopes of this volcano). 

Plants slender, 1.5-3 meters high, the stems 1.5-2.5 cm. thick, densely annu- 
late; petioles about 30 cm. long, glabrous or glabrate; leaf blades thin, somewhat 
paler beneath, about 75 cm. long, the rachis brownish-furfuraceous at first, becom- 
ing glabrate; leaf blades irregularly pinnatisect, with about 30 primary nerves on 
each side, the segments several-nerved and 1-nerved, the 2 uppermost broad and 
falcate, 15-30 cm. long and 6.5-8.5 cm. wide, long-acuminate, 2-3 pairs of the seg- 
ments several-nerved and 3-4 pairs 1-nerved, all of them conspicuously sigmoid; 
spadix about 30 cm. long, twice branched, the peduncles about 11 cm. long; scars 
of the spathes 3 cm. apart; spadix branches densely granulose and white-hirsute, 
slender, orange, the primary branches about 10, the lowest 2 divided into 2-3 
branchlets, the ultimate branchlets 7.5-10 cm. long, not spinose at the apex, the 
pits spirally arranged, the axis of the branchlets 1-1.5 mm. thick; pits with no 
upper lip, the lower lip slightly produced but truncate or scarcely rounded, entire; 
staminate flowers slightly emergent from the pits, whitish; immature fruits sub- 
globose but slightly longer than broad, about 8 mm. long. 

Geonoma longepetiolata Oerst. Vid. Medd. Kjoebenhavn 
1858: 36. 1859. 

Wet, mixed, lowland forest, at or little above sea level; Izabal. 
British Honduras, along the Atlantic coast to Costa Rica. 



268 FIELDIANA: BOTANY, VOLUME 24 

Stems 1.5-5 meters high, 2.5 cm. thick or less, the leaf scars 2-3 cm. apart; 
leaves somewhat brownish-furfuraceous when young but soon glabrate, about 1.5 
meters long, the sheath short-cylindric, 15-20 cm. long, the blades with about 30 
primary nerves on each side; leaf segments 4-8 on each side, very unequal in width, 
the lower ones much narrower, 6-12-nerved, the apical ones 10-12-nerved, 40 cm. 
long or less, sigmoid; spadix twice branched, 40 cm. long or less, the peduncle 6-8 
cm. long, the rachis 15-17 cm. long, the 12-14 branches slender; pits arranged in 
whorls in groups of 3, projecting conspicuously from the axils, the whorls 6-8 mm. 
apart, the lower lip bifid; fruit globose, 6 mm. in diameter; seed scarcely 5 mm. 
thick. 

Called "monkey-tail palm" in British Honduras; "cah-ca" in 
Honduras (an Indian name). G. flaccida WendL, reported from Gua- 
temala by Hemsley on the basis of a Skinner collection, is considered 
a synonym of G. longepetiolata by Burret, who does not, however, cite 
the Skinner collection, which may represent some other species. 

Geonoma membranacea Wendl. ex Spruce, Journ. Linn. Soc. 
11:106.1871. Cum (San Marcos). 

Moist or wet, mixed, mountain forest, 1300-1500 meters; en- 
demic; Santa Rosa (Cieneguilla) ; Escuintla (type collected by Wend- 
land on the western slope of Volcan de Fuego, between San Pedro 
and Finca Zapote); San Marcos (Volcan de Tajumulco). A photo- 
graph of the type is in the Herbarium of Chicago Natural History 
Museum (Neg. 20793). 

Stems slender, 2-4 meters high, 2.5-5 cm. thick, the leaf scars 3-5 cm. apart; 
leaves about 1.5 meters long, the sheath 10-15 cm. long, closed, the petiole 40-50 
long, the blade with 35-40 nerves on each side; segments 5-11 on each side, usually 
7-9, unequal, sigmoid, the basal ones 5-7-nerved, 2.5-5 cm. wide, 20-40 cm. long, 
the succeeding ones 1-3-nerved, the terminal segments 10-12-nerved and 8-11 cm. 
wide, the segments 2-7 cm. apart; spadix long-pedunculate, 40-50 cm. long, spread- 
ing or pendent, the spathes 12-14 cm. long, caducous; rachis of the spadix 9-17 cm. 
long, the 7-10 slender branches 15-20 cm. long; pits in 5 ranks, 7-10 mm. apart; 
fruit globose, 7-8 mm. in diameter, the globose seed 5-6 mm. thick. 

Geonoma mexicana Liebmann, ex Mart. Hist. Nat. Palm. 3: 
316. 1850. Pamac, Canpamac (Quecchi). 

Moist or wet, mixed forest, 800 meters or lower; reported from 
Pete"n; Alta Verapaz; Izabal (Sierra del Mico). Southern Mexico; 
reported to extend southward to Costa Rica along the Atlantic coast. 

Stems usually 1.5-2 meters high and 1-1.5 cm. thick, the leaf scars about 5 cm. 
apart; leaves rather small, with a very slender petiole and rachis, the segments thin, 
usually 3 pairs, broad and somewhat sigmoid, several-nerved; spadix 20-30 cm. 
long, densely pubescent, twice branched, on a short or elongate peduncle, the pits 
rather dense, only 3-6 mm. apart, the flowers conspicuously emergent; fruits glo- 
bose, 6 mm. in diameter. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 269 

Tuerckheim 8332 from Cubilgiiitz, Alta Verapaz, which is refer- 
able here, was reported from Guatemala as G. multiflora Mart., a 
South American species. Burret used for this species the name 
Geonoma oxycarpa Mart., which is a different species, of Haiti (see 
L. H. Bailey, Gentes Herb. 4: 260. 1939). Lundell has reported from 
Pete*n the names "moxan," "cambo," and "uatapil" as in use for a 
plant referred to this species. In Honduras the young inflorescences 
of the Geonomas sometimes are cooked and eaten like those of 
Chamaedorea. The leaves of various species are often used as thatch, 
at least for temporary shelters. 

Geonoma polyneura Burret, Notizbl. Bot. Gart. Berlin 11: 
500. 1932. 

Moist or wet, mountain forest, 2500 meters or lower; Alta Vera- 
paz (type from Finca Sepacuite", 0. F. Cook & R. F. Griggs 36); 
Zacapa (Sierra de las Minas). 

Caudex stout, about 3 cm. in diameter; blades apparently large, thin, slightly 
paler beneath, the segments several-nerved, falcate, narrowly long-acuminate, the 
terminal segments 10 cm. wide or more, about 40 cm. long, 15-nerved, the middle 
and lower segments narrower, as much as 55 cm. long, some of them 12-nerved; 
spadix 40 cm. long or more, twice branched, the branched portion 21-23 cm. long, 
the rachis 13 cm. long, the spathes robust, 15 cm. long or more, 2.5-3.5 cm. wide, 
the upper one inserted 2.7 cm. above the lower; primary branches of the spadix 
10-11, the lower 2-4 again branched, the branches spreading, at first fuscous- 
pilose, the ultimate branches 2-2.5 mm. in diameter, not spinose at the tip; pits 
arranged spirally in about 5 series, 12 mm. apart, bilabiate, the lower lip somewhat 
protracted, the upper one conspicuously produced; flowers about 3.5 mm. long, 
large for the genus. 

Geonoma Seleri Burret, Bot. Jahrb. 63: 211. 1930. Cum (Que- 
zaltenango) ; Kiang (San Marcos) ; Box (Huehuetenango) ; Pamaca 
(Zacapa). 

Moist or wet, mixed, mountain forest, 1300-2300 meters, or 
sometimes lower; endemic; Alta Verapaz (between Sepacuite" and 
Panzos); Zacapa (Sierra de las Minas); Quezaltenango (Volcan de 
Santa Maria) ; San Marcos (Volcan de Tajumulco) ; Huehuetenango 
(type from forests of Yalambohoch, Seler 2757; also near Maxbal). 
Figure 44. 

Plants 3-9 meters high, the caudex 2.5-7.5 cm. in diameter, conspicuously an- 
nulate; leaves large, long-petiolate, the blades unequally pinnatisect, the segments 
numerous, the terminal ones united with the rachis for 11-13 cm., 11-12-nerved, 
the lower segments of unequal breadth, mostly 3-nerved, rarely 1-nerved, as much 
as 55 cm. long; spadix twice branched, recurved, densely brownish-tomentulose at 
first, glabrate in age; upper spathe inserted 6 cm. above the lower one; peduncles 




FlG. 44. Geonoma Seleri. a, Basal portion of leaf (X V&)- b, Middle section 
of leaf (X V)- c A P ex of leaf ( x V). d, Portion of spadix (X 2 A)- , Young 
spadix and spathe (X V)- /, F 1 * (X H)- 

270 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 271 

about 20 cm. long, the primary branches of the spadix about 13, the lower ones 
branched, the ultimate branchlets spinose at the apex, as much as 3.5 mm. in diam- 
eter near the base; pits spirally arranged, about 5-ranked, 11 mm. apart or some- 
times closer together, the upper lip somewhat protracted, the margin acute, the 
lower lip emarginate; sepals of the staminate flower 4 mm. long, the petals slightly 
longer than the sepals; pistillate sepals 4 mm. long, ovate-oblong. 



MANICARIA Gaertner 
Reference: Burret, Notizbl. Bot. Gart. Berlin 11: 389. 1928. 

Plants very robust, tall or low, unarmed, the caudex stout, annulate, often 
curved or flexuous, covered with old leaf sheaths; leaves terminal, very large, sub- 
erect, lanceolate, acute, plicate-nerved, serrate at first and finally pinnatisect, the 
costa thick and stout, the petiole slender, the sheath cleft, its margins with many 
coarse fibers; spadices several, erect-spreading, tomentose, the branches strict, 
rather thick, foveolate; spathes 2, the upper fusiform, terete, mucronate, fibrous, 
tardily rupturing; bracts subulate; flowers monoecious, borne in the same spadix, 
this inserted among the leaves, simply branched, the flowers immersed in pits in 
the branches, the upper ones staminate, crowded, the lower ones scattered, pistil- 
late; staminate flowers oblong, obtusely trigonous, the sepals ovate-rounded, coria- 
ceous, with scarious margins, imbricate, the petals thick-coriaceous, obovate- 
oblong, valvate; stamens 24-30, the filaments filiform, connate at the base, the 
anthers narrowly linear, erect, bifid at the base, emarginate; pistillate flowers 
larger, ovoid, the perianth little enlarged after anthesis, the sepals rounded, their 
margins finally lacerate, broadly imbricate, the petals longer, convolute-imbricate 
at the base, acute and valvate at the apex; ovary sulcate, 3-celled, the stigmas 3, 
sessile; fruit large, globose, 1-seeded, or depressed-globose and 2-3-seeded, the 
stigmas terminal, the pericarp corticate, the cortex corky, angulate-echinate, the 
endocarp vitreous-crustaceous, fibrous within; seeds globose, erect, the hilum ob- 
long, the testa very hard, the branches of the raphe closely reticulate and involving 
the seed, the endosperm corneous, uniform, the embryo basal. 

Three species are known, the others Brazilian. 

Manicaria saccifera Gaertn. Fruct. & Sem. 2: 469. pi. 176. 1791. 
Yolillo; Confra; Manaca. Figure 45. 

Abundant in coastal swamps of the north coast, not extending in- 
land beyond the influence of tidewater. British Honduras to Brazil. 

Plants sometimes 6 meters high, but usually lower, the trunk usually short or 
almost none; leaves often 5 meters long; inflorescence about a meter long; fruit 
4-5 cm. in diameter, covered with numerous thick, irregularly pyramidal tubercles. 

This is the most abundant and conspicuous plant in the extensive 
tidal swamps along the North Coast, as at Puerto Barrios, where it 
forms wide dense stands. In Guatemala the trunks usually are short 
or the plants acaulescent, but sometimes the trunks attain here a 
height of at least 8 meters. They are much infested with mosses, 




272 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 273 

Nephrolepis (a fern), and other epiphytes. The huge leaves are much 
used along the Atlantic coast of Central America for thatch, and are 
said to last longer than those of any other palms. The soft brown 
feltlike spathes, composed of tough, closely interlaced fibers, form a 
sort of conic covering for the spadix. They are sometimes used in 
Panama for making long-peaked caps that are one of the articles sold 
to tourists there. The plant is called "gudgara" in Panama. 

OPSIANDRA 0. F. Cook 

Plants solitary, the trunk erect, ascending, or flexuous, columnar below, slightly 
tapering above, scarcely enlarged at the base, supported by a conic mass of very 
thick roots; leaves usually 5 or 6, ascending, 2-3 meters long, with a cylindric, 
sheathing base ; pinnae numerous, lanceolate, not grouped ; inflorescences inf raf oliar, 
with numerous tapering, simple branches, or a few of the lower branches furcate 
near the base; spathes 4, slender, incomplete; flowers alike, in longitudinal rows of 
2 or 3, the lowest flower of each group pistillate and somewhat smaller than the 
staminate ones, a few solitary staminate flowers present near the tips of the 
branches; sepals orbicular, broadly imbricate; petals broadly triangular, valvate, 
thick, carnose, persistent; stamens 6, the filaments short and broad; fruit globose- 
reniform, with a distinct groove on the median face above the stigma, smooth, the 
pericarp carnose; seed depressed or subreniform, almost smooth, slightly impressed 
with 5-7 simple or sparsely branched fibers radiating from the inner side of the 
hilum, the endosperm uniform, the embryo about intermediate between basal and 
lateral. 

The genus consists of a single species. It is related to Synechan- 
thus but differs in the much greater size of the plant, the pinnae 
regularly arranged but in different planes from the rachis, the stout 
rather than slender spadix branches, and the valvate rather than 
imbricate petals of the pistillate flowers. Some of the West Indian 
species of Gaussia are somewhat similar in habit. 

Opsiandra maya 0. F. Cook, Journ. Wash. Acad. Sci. 13: 182. 
1923. Palma cimarrona; Cambo. 

Known only from the forests of northern Pete"n, at 150 meters or 
lower, and in adjacent British Honduras; type from Laguna Colo- 
rada, Tikal District, Cook & Martin 94. 

Trunk 20 meters tall or more, about 15 cm. in diameter near the base, the inter- 
nodes 12-15 cm. long, becoming shorter above; leaves 2-3 meters long, the sheath- 



FIG. 45. Manicaria saccifera. a, Portion of leaf near base (X 1 A). b, Portion 
of leaf near middle (X H) c, Apex of leaf (X M)- d, Staminate flower (X 2). e, 
Portion of inflorescence, showing female flowers near base and male near apex 
( X M)- / Fruiting spathe opened to show fruits in position (X Ji). 



274 FIELDIANA: BOTANY, VOLUME 24 

ing portion 30 cm. long, the petiole 65 cm. long, 3 cm. thick above; pinnae about 
88 pairs, the lowest 41 cm. long and 2.2 cm. wide, the largest ones, somewhat below 
the middle, 61 cm. long and 3.8 cm. wide, one nerve on each side of the costa more 
prominent than the others; inflorescence 75 cm. long, the branched portion 34 cm. 
long, the branches 17-18, 30 cm. long or less, 4-5 mm. thick; spathes 4, the lowest 
9.5 cm. long, 5.5 cm. wide, distinctly carinate on each side, the uppermost 19 cm. 
long and 2 cm. wide; sepals 1 mm. long, the pistillate petals in anthesis 2 mm. long, 
slightly accrescent; fruit 1-1.5 cm. in diameter, with a soft, red, fleshy pericarp, the 
flesh mucilaginous and very sticky; seeds about 1 cm. in diameter. 



ORBIGNYA Martius 

Reference: M. Burret, Notizbl. Bot. Gart. Berlin 10: 498. 1929. 

Low or tall, unarmed palms, the trunk solitary, with irregular scars; leaves 
terminal, pinnatisect, the segments alternate, linear-lanceolate, acuminate, the 
margins recurved at the base, the sheath open; spadices large, the rachis thick, 
the branches short, alternate, erect, the pistillate thicker than the staminate ones; 
spathes several, fusiform; bracts and bractlets membranaceous, connate; stami- 
nate flowers small, the sepals minute, ovate-triangular, the petals 3, ovate or 
ovate-lanceolate, entire or dentate, free or connate at the base, valvate; stamens 
12-14, the filaments filiform, the anthers linear, spirally twisted; pistillate flowers 
large, subglobose or trigonous-ovoid, the perianth accrescent after anthesis, the 
sepals thick-coriaceous, ovate or oblong-lanceolate, imbricate, the petals similar 
but smaller, convolute-imbricate, acute, sometimes dentate; ovary ovoid, 2-7- 
celled, the style short and thick, with 2-7 large, erect stigmas, the ovules basal, 
erect; fruit large, globose or ovoid, rostrate, 2-6-seeded, the style terminal, the 
pericarp fibrous, the endocarp osseous, fibrous outside, 3-6-pored near the base; 
seeds narrow, oblong or obovoid, the testa reticulate by the branches of the raphe, 
the endosperm uniform, cartilaginous, the embryo opposite one of the pores. 

A genus of about 20 species, the others South American. 

Orbignya Cohune (Mart.) Dahlgren ex Standl. Trop. Woods 
30: 3. 1932. Attalea Cohune Mart. Hist. Nat. Palm. 3: 300. pi. 167. 
1836-50. Manaca; Corozo; Cohune; Tutz (Maya); Cords (Quecchi). 

Abundant at many places in the Atlantic lowlands, growing 
mostly on rather well-drained land, on plains or often on mountain 
sides, chiefly at 300 meters or lower; Pete"n; Alta Verapaz; Izabal. 
Southern Mexico; British Honduras to Honduras, and perhaps ex- 
tending as far south as Costa Rica. Figure 46. 

Plants very large, often low and acaulescent or with a short trunk, the trunks 
of mature trees 9-15 meters tall, thick, usually bearing persistent leaf bases above; 
leaves numerous, plume-like and graceful in spite of their great size, sometimes 
10 meters long and 2 meters wide, recurving, with numerous, elongate, very nar- 
row segments; staminate inflorescences commonly 1-1.5 meters long, with numer- 
ous flowers; fruiting panicles very large and heavy, pendent, containing often 
800-1000 fruits; fruits about 6 cm. long, resembling small coconuts. 




a 



S.H;Gr 



FIG. 46. Orbignya Cohune. a, Section of leaf ( X Vs). b, Portion of staminate 
spadix (X */*) c, Portion of fruiting branch (X Vs)- 



275 



276 FIELDIANA: BOTANY, VOLUME 24 

The names "corozo" and "manaca" are used commonly for this 
palm in Guatemala, and the term "manaca shack" is given commonly 
by the English-speaking people of the coast to the dwellings made 
from the leaves. A caserio of Izabal has received very appropriately 
the name "Manaca." 

This is by far the largest and most majestic of all the palms of 
Central America. It is abundant in many places along and near the 
Atlantic coast, often forming stands of wide extent. It is naturally 
a forest palm, forming a dense undergrowth in primeval mountain 
forest, but if the shade is removed the plants seem to thrive equally 
well, and they are seen everywhere in their native regions in open 
places. Morris, who gave considerable attention to this and other 
palms in British Honduras, estimated one leaf he saw to have a 
length of 18 meters and a breadth of 2.5 meters, and lengths of 10-12 
meters are not uncommon. The plant thus produces probably the 
largest leaves of any American plant. Brigham, who studied this 
and other Guatemalan plants many years ago, estimated the number 
of staminate flowers in an inflorescence at more than 30,000. These 
flowers attract many bees and wasps. He maintained that the plant 
was called "manaca" when acaulescent, and "corozo" when having 
a well-developed trunk, but this is questionable. 

In Guatemala the corozo often is associated with Pinus caribaea. 
It is the most conspicuous palm along the railway leaving Puerto 
Barrios, all the way inland until the dense forest terminates shortly 
before the road reaches Gualan. On the North Coast of Guatemala 
this palm is used in constructing dwellings, a very great part of them 
being made almost wholly of the materials it supplies. The stout 
midribs of the leaves form the framework, while the leaves, placed 
crosswise so that their segments form a dense thatch, constitute the 
roofs. Such buildings are substantial and durable. The young leaves 
are utilized for making hats. Segments of the large leaves are used 
by the Indians of Alta Verapaz for making the suyacales large mats 
that protect their clothing and loads from the rains. During the 
rainy season an Indian with a rolled-up suyacal (if it is not raining) 
is almost as characteristic of Alta Verapaz as the Britisher with his 
rolled umbrella is of England. 

Throughout its range, a kind of wine sometimes is made from the 
sap of the corozo. A cavity about 30 cm. square is cut in the cabbage, 
screened in various manners, and left for about a week, during which 
time the sap collects and ferments. 

The part of the corozo having the greatest possibilities is the 
seeds. These are rich in oil, which is in demand for making soap and 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 277 

for other purposes. The only obstacle to the development of an im- 
portant corozo industry is the hardness of the nuts, which are diffi- 
cult to crush. In recent years an industry of considerable importance 
has been developed in British Honduras, in cultivating the plants 
and extracting oil from the seeds, and special machinery is now used 
for their treatment. During the first world war the seeds were used 
in preparation of charcoal for gas masks. The nuts are produced in 
great abundance, as indicated in the description above, and a single 
fruiting panicle sometimes weighs more than 100 pounds. 

PAUROTIS 0. F. Cook 

References: Beccari, Webbia 2: 107. 1907; H. H. Bartlett, Car- 
negie Inst. Wash. Publ. 461: 31-34. 1935; Bailey, Gent. Herb. 4: 
361. 1940; H. E. Moore, Gent. Herb. 8: 210. 1951. 

Erect, cespitose, slender, medium-sized hermaphrodite palms, the trunk clothed 
with persistent leaf sheaths or in age naked below; leaves palmate, the sheath soft 
fibrous, unarmed; petioles armed with stoutish teeth along the margins, ligule pres- 
ent at base of the blade above, absent below; inflorescences interfoliar, erect- 
slender, with several twice-branched divisions, the tubular spathes sheathing the 
peduncle and primary branches; flowers small, in clusters of 3-2 (-1) on slightly 
elevated bracteolate cushions, the calyx of 3 nearly free sepals about half as long 
as the 3 essentially free, valvate, foveolate petals, stamens 6 with filaments united 
in a ring below, attenuate above to the anthers; carpels 3, lightly connate below, 
the styles short, connate in a conic beak and stigmatic at the apex, the ovule basal 
and erect; fruit globose or ovoid-globose, small, with smooth exocarp, slightly 
fleshy mesocarp, and thin, fragile endocarp; seed globose with a short raphe half 
as long as the seed, the endosperm homogeneous without intruded integument, 
embryo lateral. 

The genus consists of a single species and by most recent authors 
has been treated under the generic name Acoelorraphe Wendland. For 
comments on the correct name see Bailey and Moore listed above. 

Paurotis Wrightii (Griseb. & Wendl.) Britton ex Britton & 
Schafer, N. Amer. Trees 141. 1908. Copernicia Wrightii Griseb. & 
Wendl. ex Griseb. Cat. PI. Cuba 220. 1866. Acoelorraphe Wrightii 
Beccari, Webbia 2: 109. 1907. Brahea psilocalyx Burret, Notizbl. 
Bot. Gart. Berlin 11: 1037. 1934 (type from Manatee Lagoon, British 
Honduras, M. E. Peck 241)- A. pinetorum Bartlett, Carnegie Inst. 
Wash. Publ. 461: 33. 1935 (type from pine woods near Belize, British 
Honduras, H. H. Bartlett 11201}. Palma. Figure 47. 

Moist or wet pine woods, sometimes in Manicaria swamps, or 
about or in savannas, 200 meters or lower; Alta Verapaz; Izabal; 



278 



FIELDIANA: BOTANY, VOLUME 24 




(X 



FIG. 47. Paurotis Wrightii. a, Leaf (X V"). b, Portion of fruiting branch 



probably also in Pete"n. Southern Mexico to Honduras; Cuba; 
southern Florida. 

A rather small palm, the trunk slender, 3-8 meters high, 6-10 cm. in diameter, 
the plants usually forming colonies; leaf sheaths rufous-brown, glabrous, lustrous, 
dividing into numerous fibers; petioles 125 cm. long or shorter, usually coarsely 
dentate on the margins with brown incurved teeth; leaf blades grass-green above, 
dull pale green or silvery beneath, orbicular in outline, with about 40 segments on 
each side, almost glabrous or minutely whitish-pubescent; inflorescences ascending 
to erect, the spadix about a meter long, with about 9 spathes, the lowest spathe 
reddish, semicylindric, about 14 cm. long, glabrate, but when young floccose-pilose, 
especially on the margins; primary branches of the spadix 25 cm. long or less, the 
branchlets white-tomentose, only the ultimate ones floriferous, 5-12 cm. long; 
calyx segments pale brown, 1 mm. long, carnose, free and membranaceous at the 
apex, multifimbriate; corolla pale brown when dry, white-lineate, the petals fleshy, 
glabrous, 1.7 mm. long; fruit 7-8 mm. long and 6-8 mm. in diameter. 

In British Honduras called "Honduras pimenta" and "Hairy Tom 
palmetto." No good reasons have been given for separating the Brit- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 279 

ish Honduras palm from that of Cuba and Florida. Bartlett says that 
Acoelorraphe pinetorum differs from A. Wrightii "in a number of mi- 
nute but definite floral characteristics that are sufficient to give it 
specific status," but these must be minute indeed, since he gives no 
hint of what they may be. Apparently the British Honduras palm's 
chief claim to recognition as a separate species is its range; but range, 
despite its frequent invocation, is not a specific character. 

DOUBTFUL SPECIES 

Paurotis Schippii Burret, Notizbl. Bot. Gart. Berlin 12: 303. 1935. 
Brahea Schippii Burret, op. cit. 304, nomen. Acoelorraphe Schippii 
Dahlgren, Field Mus. Bot. 14: 9. 1936. 

Type from swamp forest, Pojktuun trail, British Honduras, at 
780 meters, W. A. Schipp S893. 

Plants as much as 6 meters high, the trunks solitary, 15-20 cm. in diameter; 
free portion of the petiole 70 cm. long, 1.5 cm. broad at the base, the margins com- 
pletely unarmed; ligule triangular, conspicuously produced, densely lanuginous- 
tomentose with rather long, fuscous hairs; blade 60 cm. in greatest length, white- 
ceraceous beneath, with about 60 segments, these only 24-28 cm. long, the rachis 
about 12.5 cm. long; spadix 2.5 meters long, the peduncle 1.7 meters long; spathes 
6, narrowly tubular, glabrous, striate-nerved, the lowest 36 cm. long; branched 
portion of the inflorescence 78 cm. long, the branches closely and very densely 
fuscous-tomentose, the ultimate floriferous branchlets filiform, 5-11 cm. long, the 
flowers glabrous; sepals ovate, the petals elliptic-oblong; ovary glabrous; fruit 
unknown. 

Although placed in Paurotis by Burret, this species has much 
more affinity with Erythea. Until mature fruit is known, its dis- 
position must remain in question. 

PHOENIX L. Date 

Plants unarmed, the trunk usually thick and elongate, sometimes short, often 
cespitose, covered above by the persistent leaf bases; leaves terminal, spreading- 
recurved, unequally pinnate, the segments grouped or almost equally spaced, elon- 
gate-lanceolate or ensiform, acuminate, stiff; spadices usually several, erect or in 
fruit pendent, the peduncle strongly compressed; spathe complete, elongate, com- 
pressed and 2-edged, coriaceous, cleft ventrally; flowers small, dioecious, sessile 
along the fastigiate branchlets, minutely bracteate, ebracteolate, the perianth cori- 
aceous; staminate flowers oblong or ovoid, the calyx cupular, tridentate, the 3 
petals obliquely oblong or ovate, subconnate at the base, valvate; stamens 6, the 
subulate filaments connate at the base, the anthers linear-oblong, erect, dorsifixed; 
pistillate flowers as large as the staminate, globose, the perianth accrescent after 
anthesis, the 3 petals rounded, concave, broadly imbricate; ovary 3-carpellate, the 
carpels distinct, ovoid or globose, the stigmas sessile, uncinate, the ovules erect; 



280 FIELDIANA: BOTANY, VOLUME 24 

fruit oblong, terete, 1-seeded, the stigmas terminal, the pericarp fleshy, the endo- 
carp thin-membranaceous; seed erect, linear-oblong, deeply sulcate ventrally, the 
hilum basal, the endosperm corneous, the embryo dorsal or subbasal. 

About a dozen species, in tropical and subtropical Asia and Africa, 
one of them of great importance for its edible fruit and grown for this 
reason in remote parts of the earth. 

Phoenix dactylifera L. Sp. PI. 1188. 1753. Datilero (tree). 
Datil (fruit). 

Native of northern Africa but often grown in other parts of the 
earth; planted in Guatemala infrequently, for ornament or as a curi- 
osity, but little planted anywhere in Central America. 

Although there are occasional date trees in many parts of Cen- 
tral America, most of them never produce fruits; it is not certain that 
any of them ever do. In desert areas of northwestern Mexico, and in 
southern Arizona and California, the trees thrive and fruits are pro- 
duced freely. A date industry of importance is now well established 
in the United States. Regions of similar climate are found in Guate- 
mala, particularly in the Motagua Valley and about Huehuetenango, 
and there is no apparent reason why dates should not be produced 
there, if proper treatment were given to ensure fertilization. The 
date palm was imported to Mexico by the Spaniards probably soon 
after the Conquest. In Guatemala we have observed date trees of 
thrifty appearance in Guatemala, Almolonga, Huehuetenango, Co- 
ban, and elsewhere. 

Phoenix canariensis Hort., a very common ornamental palm of 
Florida, believed to come from the Canary Islands, is planted fre- 
quently for ornament in Guatemalan gardens, especially in the cen- 
tral departments. It is somewhat similar to the date palm but much 
more slender in all its parts. 

REINHARDTIA Liebmann 

References: M. Burret, Die Palmengattungen Reinhardtia Liebm. 
und Malortiea H. Wendl. Notizbl. Bot. Gart. Berlin 11: 551-556. 
1932; H. E. Moore, Reinhardtia, Gentes Herb. 8: 541-576. 1957. 

Plants small and low, sometimes nearly acaulescent, unarmed, slender, the 
stems covered by the leaf sheaths; leaves pinnate or simple and bifid at the apex, 
the broad terminal segments of some species several-nerved with elongate perfora- 
tions or "windows" along the costa; spadices inserted between the leaves, long- 
pedunculate, erect, the branches few, strict, stiff, simple or the lower sometimes 
again branched, compressed-angulate, bracteate at the base; spathes 2, much 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 281 

shorter than the peduncle, membranaceous or coriaceous, compressed and bicari- 
nate, the flower clusters rather remote; flowers unisexual, borne on the same spadix, 
ternate on the branches, the middle flower pistillate; staminate flowers slightly 
asymmetric, conic-ovoid, the sepals orbicular, concave, imbricate; petals ovate or 
lanceolate, acute, rigid, valvate; stamens 8-40, the filaments short, the anthers 
linear, basifixed or dorsifixed; pistillate flowers smaller, ovoid, accrescent after an- 
thesis, the sepals orbicular, imbricate, petals exceeding the sepals, valvate above a 
slightly imbricate or connate base, staminodes present and variously adnate to the 
petals, the pistil trilocular, triovulate, with only one ovule usually developing; fruit 
small, ovoid or ellipsoid, apiculate, with terminal stigmatic scars, 1-celled and 
1-seeded; seed ellipsoid, erect, the attachment basal or lateral, the endosperm 
homogeneous or ruminate. 

The genus consists of five species, mostly Central American but 
extending into southern Mexico and Colombia. R. Koschnyana 
(Wendl. & Dammer) Burret of Costa Rica to Colombia is one of 
the smallest palms known. 

Seeds with deeply ruminate endosperm; stamens 27-40; petiole not pale or wing- 
margined below, densely and persistently marked with brown scales. 

R. latisecta 

Seeds with essentially homogeneous endosperm; stamens 8-22; petiole pale and 
narrowly winged along the margin below with deciduous scales . . . . R. grarilis 

Reinhardtia gracilis (Wendl.) Burret, Notizbl. Bot. Gart. Ber- 
lin 11: 554. 1932. Malortiea gracilis Wendl. in Otto & Dietr. Allg. 
Gartenz. 21 : 26, 146. 1853. Figure 48. 

Stems cane-like, solitary or cespitose in dense to loose clumps, to 2.45 meters 
high and 1.4 cm. in diameter; sheaths 6-15 cm. long; petiole elongate, pale and 
narrowly wing-margined below with deciduous scales, the rachis 3.5-23 cm. long 
with 2 or rarely 3-4 pinnae on each side, these with narrowly elongate perforations 
or "windows" along the rachis between most of the 8-22 pairs of nerves, the ter- 
minal pinnae largest, with 5-16 nerves, 7.5-23 cm. long on the upper margin, the 
outer margin coarsely dentate, the lower pair or pairs of pinnae narrowly cuneate 
and mostly 3-7-nerved, 8.5-25 cm. long, 2.8-15 cm. wide, coarsely dentate on the 
lower margin; inflorescence long-pedunculate, with 2-11 or more mostly simple 
branches 2.5-29 cm. long, becoming dark red or purplish in fruit, the staminate 
flowers 3-6 mm. long with 8-22 stamens; fruit 12-16 mm. long, 6-8 mm. in diam- 
eter when dry, black at maturity, capped with a low to prominent umbo; seed 
basally attached, the endosperm homogeneous or with minute marginal intrusions. 

According to Moore, this species consists of four varieties, dis- 
tributed from southern Mexico to Costa Rica. Two of these varieties 
are known to occur in Guatemala, R. gracilis var. gracilis, the typical 
variety, and R. gracilis var. gracilior. The latter is distinguished 
from var. gracilis in having 8-10 instead of 16-22 stamens, and in 
having staminate flowers 3-4 mm. instead of 5-6 mm. long when 
mature. 




FIG. 48. Reinhardtia gracilis. a, Habit of upper portion of fruiting plant 
(X 2 A). b, Fruit (X 3). 



282 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 283 

Reinhardtia gracilis var. gracilis 

Dense wet forests, at or little above sea level to 520 meters; Iza- 
bal; described from cultivated plants of Guatemalan origin; British 
Honduras (Pueblo Viejo, Schipp S-523} ; Honduras. 

Stems 1-2.4 meters high; leaves large with 14-22 nerves on each side of a rachis 
11-23 cm. long, the lower pinnae 4-7-nerved, 14.5-25 cm. long, 3-6.5 cm. wide; 
inflorescence with 6-11 branches, the lower often forked or branched, 7-29 cm. 
long, flowering nodes 16-40, the staminate flowers 5-6 mm. long with 17-22 sta- 
mens; fruit 14-16 mm. long with a low crown at maturity. 

Reinhardtia gracilis var. gracilior (Burret) H. E. Moore, 
Principes 1: 140, 145. 1957. Reinhardtia gracilior Burret, Notizbl. 
Bot. Gart. Berlin 11: 555. 1932. 

Moist or wet forest, at or little above sea level; British Honduras, 
the type from Stann Creek Valley, W. A. Schipp 8369; to be expected 
in Pete"n and Izabal. Southern Mexico; Honduras. 

Stems to 1 meter high; leaves small with 8-11 nerves on each side of a rachis 
3.5-6 cm. long, the lower pinnae 3-4-nerved, 8.5-12 cm. long, 1.8-3.2 cm. wide; 
inflorescence with 3-6 simple branches 2.5-8.5 cm. long, the flowering nodes 6-17, 
staminate flowers 3-3.5 mm. long with 8-10 stamens; fruit 12-14 mm. long with 
a low crown. 

Reinhardtia latisecta (Wendl.) Burret, Notizbl. Bot. Gart. 
Berlin 11: 554. 1932. Malortiea latisecta Wendl. in Otto & Dietr. 
Allg. Gartenz. 21: 146. 1853. 

Originally described from plants of Guatemalan origin, collected 
by Kegel; not collected recently in the country but to be expected in 
Izabal or the lowlands of Alta Verapaz. British Honduras (Middle- 
sex, W. A. Schipp S-56; Stann Creek Valley, Schipp 8370; Rio 
Grande, Schipp S-370). 

Plants tall, as much as 8 meters high, cespitose in clumps of 8-9 stems, these to 
6.5 cm. in diameter; leaves about 10, long-petiolate, the petiole densely and per- 
sistently brown-scaly and not wing-margined, the 2-3 broad 1 -several-nerved pin- 
nae to 62.5 cm. long, 11.5 cm. wide or more, with elongate apertures between many 
of the 21-24 nerves along the rachis, the upper pinnae broadest and 14-15-nerved; 
all the pinnae coarsely dentate along the outer margin; inflorescence exceeding the 
leaves, with 15-19 branches to 20 cm. long or more, becoming red in fruit; stami- 
nate flowers with 22-27 stamens; fruit black at maturity, ellipsoid or obovoid, 
about 17 mm. long, 11 mm. in diameter; seed laterally attached with deeply rumi- 
nate endosperm. 

Rhapis humilis Blume, probably a native of China, is cultivated 
rarely in Guatemala for ornament. It is a handsome dwarf palm, 



284 FIELDIANA: BOTANY, VOLUME 24 

generally 2 meters high or less, forming dense clumps of stems. The 
slender stems are green, reedlike, and unarmed; the small leaves are 
fan-shaped and 20 cm. long or shorter. 

ROYSTONEA O. F. Cook 

Tall, unarmed palms, the caudex solitary, more or less swollen at or above the 
middle, green above and almost smooth; leaves terminal, large, equally pinnatisect, 
the numerous segments narrowly linear-lanceolate, narrowed at the apex and un- 
equally bifid, paleaceous beneath, the sheath elongate; spadix large, the branches 
elongate, slender, pendulous; spathes 2, complete, the lower semicylindric, equaling 
the spadix, the upper ensiform, ventrally cleft; bracts and bractlets scale-like; 
flowers small, white, monoecious in the same spadix, arranged in lax spirals, the 
lower flowers ternate, the middle one staminate, the upper flowers solitary or bi- 
nate, staminate, the spadices arising below the leaves; staminate sepals minute, 
broadly ovate, scarious, imbricate at the base, the petals subequal, ovate, acute, 
connate at the base, thin-coriaceous, valvate; stamens 6, 9, or 12, exserted, the 
filaments short, broadly subulate, united at the base with the petals, the anthers 
large, ovate-sagittate, dorsifixed, versatile; pistillate flowers smaller than the stami- 
nate, the perianth scarcely enlarged after anthesis, the sepals ovate-rounded or 
reniform-rounded, broadly imbricate, the corolla urceolate, trilobate to the middle, 
the triangular lobes acute, valvate; ovary subglobose, gibbous at the base, 2-celled, 
the stigmas thick, sessile, the ovule parietal, ascending; fruit small, obovoid or 
oblong-obovoid, curved, 1-celled, the stigmas subbasal, the pericarp fleshy and 
fibrous, the endocarp thin; seed oblong-reniform, the testa adnate to the endo- 
carp, the hilum rather broad, ventral, the branches of the raphe radiating from the 
hilum and reticulate, the endosperm uniform, the embryo subbasal. 

Six species are now recognized, ranging from southern Florida 
through the West Indies to northern South America, and along the 
northern Atlantic coast of Central America. The name Oreodoxa 
usually has been employed for this genus, but it has been found to 
apply properly to a South American palm of different relationship. 

Fruit obo void-oblong, longer than broad; leaf sheaths not lepidote; leaves hori- 
zontal R. oleracea 

Fruit ovoid, about as broad as long; leaf sheaths brown-lepidote; leaves somewhat 
drooping R. regia 

Roystonea oleracea (Jacq.) 0. F. Cook, Bull. Torrey Club 28: 
554. 1901. Areca oleracea Jacq. Sel. Stirp. Amer. 1 : 278. pi. 1 70. 1763. 
Oreodoxa oleracea Mart. Hist. Nat. Palm. 3: 166. pi. 156, 163. 1837. 

Not seen wild by the writers in Guatemala but to be expected 
there; it or a closely related species is found wild in the lowlands of 
Honduras and British Honduras and is certainly to be expected on 
the North Coast of Guatemala. West Indies. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 285 

Plants very tall, sometimes 40 meters high, the trunk slender, somewhat en- 
larged at the base, sometimes also at or above the middle but in age becoming more 
uniform, pale; leaves 3-7 meters long and 2 meters wide, the pinnae 100 or more, 
closely set, as much as 5 cm. wide, in a single row on each side of the rachis; in- 
florescences very large and pendent, twice branched, the branches somewhat undu- 
late; fruits oblong and usually somewhat curved, 1.5-2 cm. long, 1 cm. thick, 
purplish to black. 

Called "yagua" in Honduras. 

Roystonea regia (HBK.) 0. F. Cook, Science II. 12: 479. 1900. 
Oreodoxa regia HBK. Nov. Gen. & Sp. 1: 244. 1815. Palma real. 
Royal Palm. 

Native of Cuba; now grown for ornament in most parts of trop- 
ical America; planted abundantly in Guatemala, especially in parks. 

A tall palm, sometimes 25 meters high but generally lower, the trunk swollen 
at or near the middle and commonly also near the base; leaves shorter than in the 
preceding species, the pinnae evidently in 2 rows on each side of the rachis, 3-4 cm. 
wide; branches of the spadix not undulate; fruit elongate-globose or almost globose, 
somewhat narrowed at the base, 8-13 mm. long, about 10 mm. broad, reddish 
brown or purplish at maturity. 

This is the most highly esteemed of the cultivated ornamental 
palms of Guatemala. It is planted freely in many of the parks, from 
the capital down to the coasts, also in gardens and fincas. It is much 
more common than R. oleracea. Some of the driveways leading into 
the coffee fincas of the Pacific boca costa are bordered by handsome 
avenues of royal palms. Trees of what appear to be both the species 
listed here may sometimes be seen in the same avenue. 



SABAL Adanson 

References: Beccari, Webbia 2: 10. 1907; H. H. Bartlett, Carnegie 
Inst. Wash. Publ. 461: 35. 1935; L. H. Bailey, American palmettoes, 
Gentes Herb. 3: 274-339. /. 145-190. 1934; Revision of the American 
palmettoes, Gentes Herb. 6: 366-459. /. 186-459. 1944. 

Plants tall or low, unarmed, the trunk usually thick, annulate, covered above 
by remains of the leaf sheaths; leaves terminal, orbicular, fan-shaped, flabellately 
multifid, the segments linear, bifid, often with threadlike fibers along the margins, 
the rachis short or elongate, the ligule short, adnate to the rachis, the sheath short; 
flowers perfect, the spadices large, branched, at first erect, the slender branches 
decurved or pendulous; spathes long-sheathing, tubular, the opening oblique, the 
bracts and bractlets minute, the flowers small, glabrous, solitary and sessile along 
the branches, the perianth not changed after anthesis; calyx cupular, unequally 
tridentate, truncate at the base; corolla deeply trilobate, the lobes ovate-oblong, 
acute, concave, the margins imbricate below the valvate apices; stamens 6, the 



286 FIELDIANA: BOTANY, VOLUME 24 

filaments subulate, dilated at the base and connate with the corolla tube to form 
an annulus, the anthers ovate-cordate, dorsifixed; ovary trigonous, 3-celled, atten- 
uate to a columnar style, the stigma truncate, the ovules basal, erect; fruit small 
or medium-sized, usually of a single 1-seeded carpel, the style basal, the pericarp 
fleshy, separating from the membranaceous epicarp; seed erect, depressed-globose, 
the small hilum basal, the endosperm undivided, the embryo dorsal. 

About 25 species, ranging from southern United States and Mex- 
ico to West Indies and Venezuela. No other species are known from 
Central America. 

Ligule of the leaf very small and narrow or sharp-pointed, usually not more than 

2-3 cm. long S. mayarum 

Ligule of the leaf elongate or broad or large, 4-5 cm. long or larger. 

Fruits 13-20 mm. broad, 15 mm. long; undivided basal portion of the leaf one- 
third to one-fourth the length of the blade S. mexicana 

Fruits 8-10 mm. broad, 10-13 mm. long; undivided portion of the leaf very 

short, sometimes almost none. 
Branchlets (rachillae) of the inflorescence very slender, 1 mm. thick or less 

when dry, the ends almost filiform S. nematoclada 

Branchlets relatively stout, not filiform, more than 1 mm. thick when dry. 

S. Morrisiana 

Sabal mayarum Bartlett, Carnegie Inst. Wash. Publ. 461: 35. 
1935. 

British Honduras, at or little above sea level; type from Maskall, 
Belize District, P. E. Gentle 1156; also in Corozal District; to be ex- 
pected in Pete'n. 

Plants rather small, 4 meters high; leaves about 2 meters wide, light green but 
not glaucous, somewhat paler beneath, cleft nearly to the base into many narrow 
segments, the basal undivided portion 15-20 cm. long or shorter; petioles about 
2 cm. broad at the apex, continued into the blade for 60 cm. as a rachis or costa; 
ligule very short, triangular, acute or acuminate, 2-3 cm. long; spadix 1 meter long 
or more, much branched, the sheaths finely striate, the branchlets 3-6 cm. long, 
slender, sharply angulate when dry; flowers 3-4 mm. long, contiguous but not con- 
gested on the rachilla, subtended by 3 unequal bractlets; calyx 1.5 mm. high, not 
costate, the 3 lobes obtuse, spreading; petals flat, open at anthesis, the corolla 
4-5 mm. broad when dry; fruit unknown. 

The species was reported by Bartlett from Yucatan and Cuba 
but according to Bailey, the collections so reported belong to other 
species not found in either Guatemala or British Honduras. Known 
in British Honduras by the names "botan," "huano," "bayleaf 
palm," and "big thatch." 

Sabal mexicana Martius, Hist. Nat. Palm. 3: 246. 1838. ?S. 
guatemalensis Beccari, Webbia 2: 68. 1907 (type collected in Guate- 
mala by Skinner, the locality unknown). 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 287 

Open, rather dry hillsides, or on coastal plains or in river valleys, 
sometimes along the borders of mangrove or bamboo swamps, 1400 
meters or lower; reported from Pete"n, perhaps in error; El Progreso; 
Zacapa; Chiquimula; Escuintla; Retalhuleu; San Marcos; Huehue- 
tenango(?); probably in all the Pacific coast departments. Widely 
distributed in Mexico. Figure 49. 

Younger plants often large but acaulescent, the older ones with a trunk 10-12 
meters high terminating in a crown of large and handsome leaves, often fruiting 
when acaulescent or nearly so; trunk coarsely ringed with separate steplike pro- 
tuberances, often bearing below the leaves projecting old petioles; leaf blades about 
a meter long and broad, or sometimes smaller, the main segments 4-6 cm. wide; 
petioles slender, equaling or exceeding the blade, the ligule 10-13 cm. long; inflo- 
rescences equaling or slightly exceeding the leaves; branchlets glabrous, slender, 
12 cm. long or shorter, closely flowered; flowers almost white, fragrant, 3-4 mm. 
long, with spreading petals; calyx cupular, notched, somewhat costulate; petals 
narrowly oblong, acute, nerved, longer than the calyx and equaling the anthers; 
fruits often very numerous and crowded, irregularly rounded-oblate, flattened at 
the base, 13-20 mm. broad, 15 mm. high; seed flat at the base, convex on the upper 
surface, chocolate-brown, 9 mm. in diameter; micropyle centrally lateral. 

This palm is conspicuous along the Motagua Valley in El Pro- 
greso and Zacapa, usually growing in the shallow stream beds and 
forming large groves. It is abundant at many places along the Pa- 
cific plains, sometimes forming very large groves or forests of widely 
or closely spaced trees, and forming a conspicuous feature of the 
landscape. It grows there also in cattle pastures and in remnants 
of former woodlands. The tree is often planted for ornament in 
Guatemala, and sometimes probably for its leaves, which are used 
as thatch and for the innumerable purposes for which palm leaves 
are utilized in Guatemala. Lundell states that the palm is planted 
in Pete*n, and the leaves used there for making hats, its Pete"n name 
being "guano" or "huano de sombrero." In the Motagua Valley and 
in Chiquimula the leaves are much used for thatch, and also over- 
lapped to make the sides of huts, which in that torrid region need 
not be very substantial. Palms probably of this species were ob- 
served in cultivation in Puerto Barrios. Bailey considered Sabal 
guatemalensis a name of questionable identity, but suggested that it 
may be synonymous with S. mexicana. This seems probable, since 
it is improbable that any Sabal distinct from that occurs in any part 
of Guatemala through which Skinner traveled, all the areas known 
to him probably having been covered by our own collecting trips. 

Sabal Morrisiana Bartlett, Carnegie Inst. Wash. Publ. 461: 22, 
/. 1, pi. 3, 6. 1935, without description; L. H. Bailey, Gentes Herb. 6: 
412. /. 217, 218. 1944. Botan. 




a 



FIG. 49. Sabal mexicana. a, Portion of leaf (X M)- b, Portion of fruiting 
branch (X M)- c, Habit (greatly reduced), d, Fruit, showing calyx (X 
e, Flower (X 5). 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 289 

In forest, little above sea level, Pete"n (type from Uaxactun, 
H. H. Bartlett 12284). British Honduras (Stann Creek Valley and 
elsewhere) . 

Plants very slender and tall, sometimes 27 meters high, the trunk 20-25 cm. 
in diameter, smooth, light-colored, the wood hard; leaves 2 meters broad or less, 
cleft almost to the base, light green above, silvery beneath, the segments 4-5 cm. 
wide toward the base, long-tapering; petiole equaling the blade or slightly longer, 
somewhat scurfy, the ligule 7-10 cm. long, long-acuminate; inflorescences conspic- 
uously exceeding the leaves, divaricately branched, the branchlets stout, terete, at 
least 1 mm. thick when dry, not filiform at the ends, 5-7 cm. long; fruit black, lus- 
trous, globose, somewhat narrowed at the base, 9-10 mm. broad, 10-11 mm. high; 
seed lustrous dark brown, 7-8 mm. in diameter, the micropyle lateral and prominent. 

In Pete"n the leaves are highly prized as thatch, and on this ac- 
count the trees usually are left when land is cleared for cultivation. 

Sabal nematoclada Burret, Repert. Sp. Nov. 48: 256. 1940. 

In open or rather dense forest, sometimes growing with pines and 
on limestone, at or little above sea level; British Honduras (type from 
Forest Home, W. A. Schipp 844$; San Agustin and elsewhere). 

Plants very tall and slender, as much as 27 meters high; leaves about 2 meters 
long, green above, somewhat paler but not silvery beneath, very deeply divided 
into numerous segments, these 6 cm. wide; ligule of the leaf 6-7 cm. long, narrowly 
acuminate; spadix much branched, the branchlets very slender, 1 mm. thick, al- 
most filiform at the apex, 12 cm. long or shorter; flowers cream-colored, closely 
placed, 3 mm. long, subtended by ovate-acuminate bracts and bractlets; calyx 
narrowly obovate-oblong, not conspicuously nerved, the lobes ovate-triangular; 
petals when dry narrow and with involute margins; fruit black or nearly so, lus- 
trous, obovoid-pyriform, 8-10 mm. in diameter; seed about 8 mm. in diameter, 
free from the mesocarp, the micropyle supralateral, obscure. 



SCHEELEA Karsten 

Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 651. 1929. 

Low or tall, unarmed palms, the caudex solitary; leaves terminal, pinnatisect, 
the segments seriate or grouped, linear, 1-nerved, the sheath short, open; spadices 
on long or short peduncles, the branches rather short, suberect; spathes 2, the upper 
fusiform, ligneous, acuminate, the bracts minute; flowers dioecious or monoecious 
on the same spadix, the staminate very numerous, sessile on the upper part of the 
branches, crowded, the pistillate few and solitary on the lower part of the branch, 
sessile or pedunculate; spadices inserted among the leaves; staminate sepals minute, 
triangular, acute, the petals much longer, elongate-claviform or cylindraceous, 
fleshy, acute, valvate; stamens 6, much shorter than the petals, the filaments fili- 
form, unequal, the anthers linear, affixed by the bifid base, erect; pistillate flowers 
much larger than the staminate, ovoid, the perianth greatly enlarged after anthe- 
sis, the sepals large, coriaceous, ovate, nerved, imbricate, the petals similar but 



290 FIELDIANA: BOTANY, VOLUME 24 

somewhat larger, convolute-imbricate; ovary 3-celled, the style short, terete, the 
stigmas large, spreading, the ovules basal, erect; fruit large, ovoid or oblong, 1-3- 
seeded, the style terminal, the pericarp fibrous, the endocarp thick, osseous, fibrous 
outside, 3-pored above the base; seeds ellipsoid, the testa veined by the branches of 
the raphe, the endosperm corneous, uniform, the embryo opposite one of the pores. 

The genus consists of about 40 species, most of them South Amer- 
ican. Three or four besides those enumerated here are known from 
Mexico and Central America. 

Fruiting perianth 30-35 mm. long; plant of the Atlantic lowlands S. Lundellii 

Fruiting perianth 12 mm. long; plants of the Pacific plains S. Preussii 

Scheelea Lundellii Bartlett, Carnegie Inst. Wash. Publ. 461: 
46. pi. 1-5. 1935. Corozo; Cantutz (Maya). 

In forest, little above sea level (250 meters or less) ; Pete"n (type 
collected near Polol, C. L. Lundell 3752; reported from Rio Subin and 
Rio Pasion) ; Alta Verapaz (between Candelaria and Samanzana) . 

Mature plants as much as 20 meters high, the trunk naked below, covered 
above by the remains of old leaves and inflorescences; leaves up to 8 meters long, 
the lowest ones almost horizontal, the petiole quadrangular, dilated at the base, 
splitting at the base into long, stiff, flexuous fibers, the rachis quadrangular, densely 
covered beneath with a rufous-ferruginous indument; leaf segments numerous, 
rigid, entire, the lowest 90-120 cm. long, 1.4-2.8 cm. wide, the middle ones 120- 
125 cm. long and 6-6.5 cm. wide, irregularly grouped and spaced; upper spathe 
ligneous, the principal portion fusiform, 90 cm. long, 25 cm. in diameter, attenuate 
into a spinelike, terminal appendage 45 cm. long; lower spathe 80 cm. long; pedun- 
cle ferruginous-furfuraceous, the spadix densely and simply branched, the branches 
25-35 cm. long, 2-4 mm. thick; staminate flowers 16-18 mm. long, the calyx seg- 
ments only 1.5 mm. long, the petals clavate; anthers 3.5-4 mm. long; fruit ellip- 
soid, rostrate at the apex, 6.5 cm. long, 2.8-3.4 cm. in diameter, dark brown, the 
endocarp osseous, almost 5 cm. long. 

In general appearance this and the following species are almost 
exactly like the common cohune, Orbignya Cohune. In flower struc- 
ture the two genera are unlike, the anthers of Orbignya being greatly 
elongate and spirally twisted, those of Scheelea short and straight. 
It is stated that in Pete"n Scheelea Lundellii and Orbignya Cohune 
grow together, forming forests or corozales characteristic of that region. 

Scheelea Preussii Burret, Notizbl. Bot. Gart. Berlin 10: 678. 
1929. Corozo; Coquito; Manaca. 

Abundant at many places on the Pacific plains, and probably in 
all the coastal departments, at or little above sea level, ascending to 
perhaps 900 meters but most common at lower elevations; Escuintla; 
Suchitepe*quez ; Retalhuleu; San Marcos; perhaps endemic; type col- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 291 

lected by Preuss in Guatemala, on the "Pacific side," but the locality 
unknown. Southern Mexico (Chiapas). Figure 50. 

Plants large and massive, in appearance like Orbignya Cohune, large plants 
often acaulescent, but the trunk elongating in age and commonly 3-6 meters high 
or taller, thick and very solid; leaves large, 3.5 meters long or often considerably 
longer, the rachis narrowly triangular in cross section, fuscous-leprous dorsally, the 
segments narrowly reduplicate at the base, equally spaced and inserted in the same 
plane, the larger ones 120 cm. long, the terminal segments about 30 cm. long; spathe 
about 140 cm. long, cymbiform (when opened), ligneous, slender and scarcely more 
than 5 cm. in diameter; spadix about 115 cm. long, the branched portion 70 cm. 
long, the branches numerous, 30 cm. long, almost filiform at the apex; staminate 
flowers 15 mm. long; fruits very numerous, elongate-ellipsoid, about 5 cm. long and 
2.5-3 cm. broad, rounded at each end, rostrate at the apex, the beak 3-5 mm. long; 
pistillate sepals ovate-oblong, the petals of equal length, rounded. 

This is probably a forest palm by nature, but it is most plentiful 
now on open pastured plains from which the original forest has been 
cut. In many places along the Pacific plains it is abundant, forming 
wide stands almost palm forests. Much of the land on which it is 
found is very wet during the invierno, and some of the trees stand in 
depressions that are wet throughout the year. We have not seen it 
in Santa Rosa but there is in that department an aldea called El 
Corozal, which probably attests its presence there, and it is rather 
probable that this species may extend into western El Salvador. 
The names "corozo" and "manaca" are both given commonly to this 
palm along the Pacific coast, just as they are applied to the so similar 
Orbignya of the Atlantic lowlands. 

This palm is of great economic importance locally. Its leaves are 
the commonest thatch of the humbler dwellings of the Pacific coast, 
and even of some of the more substantial ones. San Sebastian Retal- 
huleu is celebrated for its manufacture from the corozo of suyacales 
(rain capes), which are sold locally and exported to distant parts of 
Guatemala, also for its sopladores, the fans so necessary for starting 
kitchen fires, particularly those burning charcoal. Bunches of the 
nerves from the leaves are used as brooms and coarse brushes. The 
cabbage, called "palmito," is eaten either raw or cooked. Sap that 
collects in the cavity from which the cabbage is cut is allowed (sur- 
reptitiously, since it is forbidden by law) to ferment with panela or 
crude brown sugar to form an intoxicating beverage like that pre- 
pared from Acrocomia. The fruits and nuts are eaten by cattle, and 
the scant flesh is often eaten by people. The kernels are said to be 
used sometimes for making confections, and there is extracted from 
them an oil used in making soap. At San Antonio Suchitepe"quez 
there is a small factory where the oil is expressed by pressure; it is 




292 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 293 

used to make the Jabon Vegetal that is highly esteemed in Guate- 
mala for toilet purposes. The nuts are cut transversely and shaped 
to make small pipe bowls. The trunks are used for various kinds of 
construction and sometimes for fuel. 

SGHIPPIA Burret 

Plants unarmed, the caudex cylindric, without spinose roots; petioles unarmed, 
biconvex; leaf blades fan-shaped, many-cleft, without tomentum, the segments not 
filiferous; rachis none; spathes broad, the lowest sterile, the succeeding ones sub- 
tending twice-branched inflorescences; flowers spirally arranged on the branchlets, 
solitary, perfect and staminate, pseudopedicellate; calyx small, the 3 sepals free, 
lance-triangular; corolla much longer than the calyx, the 3 petals free, oblong, re- 
flexed; stamens 6, hypogynous, equal, the filaments free, elongate; anthers dorsi- 
fixed, versatile, linear, deeply sagittate at the base; carpel of the ovary 1, ovoid, 
1-celled; style narrowly tubular, obliquely expanded above into the stigma; ovule 
basal, ascending; fruit rather large, globose, the persistent style base apical, the 
perianth scarcely accrescent in age, the pericarp thin, fragile, fleshy within; seed 
globose, with a basilar hilum, the raphe basilar, not conspicuous; endosperm corne- 
ous, uniform, slightly excavate at the base, the embryo lateral. 

The genus was named for William A. Schipp, who made what is 
probably the most important collection of plants ever gathered in the 
colony of Belize, one from which a very large number of new species, 
many of them important ones, have been described. He seems to 
have had, more than any other collector who has worked in British 
Honduras, the faculty of finding and collecting the interesting plants 
of the region, rather than the common and weedy ones. Burret con- 
siders Schippia most closely related to the South American genus 
Tessmanniophoenix. 

Schippia concolor Burret, Notizbl. Bot. Gart. Berlin 11: 868. 
1933. 

Moist or wet, lowland forest, mostly at 200 meters or lower; 
Pete'n (west of Chinaja, Steyermark 45538}. British Honduras, the 
type from 19 Mile, Stann Creek Valley, W. A. Schipp S367. 

Plants about 10 meters high, the trunk 10 cm. in diameter; petiole about 2 
meters long, fuscous-tomentose below, glabrate above, 1 cm. broad, the ligule tri- 



FIG. 50. Scheelea Preussii. a, Basal part of leaf, showing basal portions of 
leaf segments (X H)- & Middle part of leaf rachis, showing basal portions of 
leaf segments ( X M)- c> Apex of leaf, with basal portions of leaf segments ( X H)- 
d, Habit of tree (greatly reduced), e, Pistillate flower (X %). /, Spathe (X '/) 
g, Portion of staminate inflorescence (X ^). h, Portion of pistillate inflorescence 
(X H). 



294 FIELDIANA: BOTANY, VOLUME 24 

angular, acuminate; leaf blade almost concolorous, 65 cm. long, cleft about two- 
thirds its length, with about 32 segments, these 3-3.5 cm. wide, gradually atten- 
uate upward; spadix twice or thrice branched, pendent, about 60 cm. long; sterile 
spathes about 25 cm. long, cochleariform, contracted at the apex, floccose-tomen- 
tose outside with a yellowish tomentum, the fertile spathes about 4, the uppermost 
8.5 cm. long, the partial inflorescences large, the 2 lowest 25 cm. long or more, the 
upper ones smaller; branchlets numerous, regularly inserted, often approximate, 
slender, glabrous, the larger ones about 12 cm. long; flowers inserted spirally, 
solitary, on pedicels 3-4 mm. long; sepals narrow, almost linear-triangular, scarcely 
1 mm. long; petals reflexed, membranaceous, 3 mm. long, oblong, subcucullate at 
the apex; stamens 3 mm. long, the filaments slightly dilated below and applanate; 
anthers linear, 1 mm. long; ovary ovoid, glabrous, abruptly contracted into the 
style; fruit globose, about 2.5 cm. in diameter when dried, lustrous, minutely gran- 
ulose; seed subglobose or depressed-globose, rounded at the apex, the largest 18 
mm. high and 20-22 mm. broad. 

Called "silver pimento" and "mountain pimento" in British 
Honduras. 

SYNEGHANTHUS Wendland 

Unarmed plants, often gregarious, the stems slender, annulate, often stolo- 
niferous; leaves terminal, equally pinnatisect, the segments broad or narrow, thin, 
plicate-nerved, acuminate, often interrupted, the petiole canaliculate above, the 
sheath short, open; spadices several, long-pedunculate, erect in flower, the branches 
very numerous and slender, flexuous, strict, compressed; spathes several, tubular, 
membranaceous, persistent; flowers minute, monoecious in the same spadix, ar- 
ranged in alternate, elongate, 1-2-seriate groups, the upper flowers of the group 
staminate, the lowest one pistillate, ebracteate and ebracteolate; staminate flowers 
trigonous-hemispheric, the calyx cupular, fleshy, trilobate; petals 3, connate at the 
base, concave, spreading, valvate; stamens 3 or 6, the filaments short, subulate, 
the anthers globose-didymous, dorsifixed; staminate flowers about as large as the 
pistillate, hemispheric, the perianth unchanged after anthesis; petals 3, orbicular, 
imbricate; ovary globose, 3-celled, the stigmas minute, sessile, the ovules erect; 
fruit ellipsoid, 1-seeded, the stigmas basal, the pericarp with thin flesh and numer- 
ous fibers, the endocarp membranaceous; seed erect, free, ellipsoid, the hilum small, 
basal, the raphe sigmoid, the embryo infra-apical. 

The genus consists of six species, four in Central America, one in 
Mexico, and one in Ecuador. In appearance the plants are similar 
to Chamaedorea, but they differ in the inflorescence, whose long slen- 
der flexuous bunched branches give it a broom-like appearance. 

Synechanthus fibrosus Wendland, Bot. Zeit. 16: 145. 1858; 
Bot. Mag. pi. 6572. 1881. Chamaedorea fibrosa Wendl. Index Palm. 
12, 57. 1854; Collinia fibrosa Wendl. loc. cit.; Rathea fibrosa Karst. in 
Koch & Fint. Wochenschr. 1 : 377. 1858. Pacaya; Capuca; Coquilla.. 

Wet forests, chiefly at or near sea level, but ascending to about 
900 meters; often growing on limestone; originally described from 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 295 

cultivated plants of Guatemalan origin; Alta Verapaz; Izabal. Brit- 
ish Honduras; Honduras. Figure 51. 

Plants slender, rarely as much as 6 meters high, usually much lower and often 
flowering when no more than 1.5 meters high, the stems smooth, green, cane-like; 
leaves few, green on both surfaces, about a meter long but often larger, the seg- 
ments linear to lanceolate, numerous, mostly 1.5-3 cm. wide, or a few of the upper 
ones much wider; spadices inserted among or just below the leaves, the peduncles 
greatly elongate; branched portion of the spadix 30-40 cm. long, the branches usu- 
ally very numerous but sometimes few, pendent in age; flowers scarcely more than 
1 mm. long, the sepals strongly costate-nerved; fruit oblong or ellipsoid, becoming 
yellow and at maturity cherry-red, as much as 2 cm. long. 

Called "monkey-tail palm" in British Honduras. 

THRINAX L. f. 

Reference: L. H. Bailey, Thrinax the peaberry palms, Gentes 
Herb. 4: 128-149. /. 75-55. 1938. 

Rather small palms, unarmed, the trunk naked, more or less annulate; leaves 
fan-shaped, long-pedunculate; inflorescences inserted between the leaves, the flow- 
ers perfect, minute; leaves divided for half their length or more, the petiole smooth, 
the ligule with a point or rim projecting over the blade; spadix usually equaling or 
exceeding the petioles, branched, glabrous or partly so, provided with small pointed 
secondary spathes at the nodes; flowers very small, consisting of a cup with 6 short 
lobes in a single series and no petals, the calyx persistent beneath the fruit; sta- 
mens 6, the filaments more or less expanded at the base and sometimes mistaken 
for petals after the anthers fall, the anthers long, exserted, 2-celled, introrse, often 
reflexed in an thesis; ovary 1-celled, globose or short-oblong, narrowed into a colum- 
nar style; fruit globose, bearing the persistent style, the exocarp succulent at full 
maturity but soon drying hard, white to gray or purplish when fresh but becoming 
blackish or brown when dry, glabrous at maturity, 10-12 mm. or less in diameter; 
seed free, the raphe obscure, the embryo and micropyle lateral toward the apex. 

Ten species were recognized by Bailey, most of them West Indian, 
but the genus is represented also in southern Florida, the Yucatan 
Peninsula of Mexico, and northern south America. No other species 
have been found in Central America, but the genus may occur along 
the Atlantic coast of Honduras. 

Thrinax parviflora Swartz, Fl. Ind. Occ. 1: 614. pi. 13. 1797. 
T. Wendlandiana Beccari, Webbia 2: 265. 1907. 

At sea level, often on coral rocks or sand; British Honduras (All 
Pines, and reported elsewhere); Turneffe Island; Florida Keys; Ja- 
maica; Haiti; Yucatan Peninsula of Mexico; Honduras(?). Figure 52. 

A slender erect palm 3-9 meters high, the trunk enlarged at the base, at the 
middle sometimes 10 cm. in diameter and obscurely if at all annulate; leaves light 




FIG. 51. Synechanthus fibrosus. a, Portion of leaf (X K). & Lower portion 
of flowering branch ( X K)- c> Lower portion of fruiting branch ( X J^). d, Habit 
of fruiting plant (greatly reduced). 



296 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 297 

green, the blades sometimes 1 meter wide or more, the petioles 60-100 cm. long, 
thin and 2-edged, bearing many marginal fibers below; ligule broad, with a low 
upturned margin, produced into a prominent triangular point; leaf blades cleft 
almost to the middle into 50 or more segments, these mostly narrow, sometimes 
3 cm. wide; spadix about a meter long, the ultimate branchlets 4-8 cm. long, the 
flowers rather closely placed, conspicuously pedicellate, variable in size; anthers 
longer than the filaments, the stigma expanded as if urceolate; fruit globose, 6 mm. 
in diameter, drying regularly with a thin brittle shell, this finely granulate; seed 
4-5 mm. in diameter. 

In Yucatan this palm is said to be known by the Maya name 
"chit." Thrinax microcarpa Sargent was said by Bailey to be "sup- 
posedly in Yucatan and British Honduras," but we know of no evi- 
dence that it grows there. 



WASHINGTONIA Wendland 
Reference: L. H. Bailey, Gentes Herb. 4: 53. 1936. 

Plants, when well developed, tall and robust, the trunk solitary, columnar, 
covered above or sometimes throughout with pendent remains of dead leaves; 
leaves large, filiferous, glabrous or nearly so, the petioles stout, compressed, armed 
with spines on the margins, the blades rounded and fan-shaped; spadices inserted 
among the leaves, 2-4 meters long, usually projecting beyond the leaves and re- 
curved or pendent, 1-3 times branched, the branchlets slender, glabrous; principal 
spathe about a meter long, compressed and 2-edged, 10 cm. broad; flowers perfect, 
whitish, short-pedicellate, about 8 mm. long (including the stamens), the calyx 
tubular, almost equaling the 3 strongly reflexed, narrow corolla lobes, persistent; 
stamens 6, attached at the base of the corolla, the anthers large and elongate, ver- 
satile; ovary trilobate, 3-celled, the style long, straight, tridentate at the apex; 
fruit hard, oblong to oblong-globose, 5-10 mm. long, smooth, with scant flesh, 
brown; seed brown, the endosperm white, hard, uniform. 

Two species are known, native in southern California and Arizona 
and northwestern Mexico, where they grow in dry, semi-desert re- 
gions. They have become favorite palms in cultivation not only in 
America but in other regions, partly because of their hardiness and 
also because they thrive under adverse conditions, either out of doors 
or as house plants. 

Washingtonia filifera (Lind.) Wendl. Bot. Zeit. 37. 68. 1879. 
Pritchardia filifera Lind. Cat. 1876, ex Andre", 111. Hort. 24. 1877. 
Palma peluda. 

Native of southwestern Arizona, southeastern California, and 
northern Sonora and Baja California; often planted for ornament in 
Guatemala, especially in the central region, but also in other places, 
such as Zacapa, Retalhuleu, and Almolonga; plants often seen grow- 




FIG. 52. Thrinax parviflora. a, Portion of leaf (X Vio)- b, Flower (X 5). 
c, Flowering branch and attachment to stem (X /^). d, Fruiting branch and 
attachment to stem (X }/%). 



298 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 299 

ing in places where the seeds probably were dispersed by accident, 
and not purposefully planted. 

Trunk sometimes 15 meters high and 1 meter in diameter, but in cultivation 
usually smaller, often covered throughout with reflexed dead leaves, but these more 
often removed by man or destroyed by fire; petiole 2 meters long or less and 10-15 
cm. broad at the base, the somewhat uncinate marginal spines 1 cm. long or less; 
blades as much as 2 meters in diameter; spadices mostly 3-4 meters long at ma- 
turity, pendent in fruit; calyx 5 mm. long, shallowly lobate, the striate petals 
reflexed to the base of the flower, the stamens about twice as long as the calyx; 
fruit short-oblong to ovoid, 7-10 mm. long; seed 5-7 mm. long. 

GYGLANTHACEAE 

Large and coarse, acaulescent, perennial herbs, palm-like in appearance, or 
scandent shrubs with annulate vaginate rooting stems; leaves distichous or spirally 
arranged, petiolate and flabellate, entire, bifid, or biparted, parallel-nerved; flowers 
monoecious, those of each sex densely crowded in whorls or spirals on a thick 
spadix; spadices axillary, solitary, pedunculate, simple, the peduncle short or elon- 
gate, subtended at the base by a bicarinate cataphyll; spathes 2-6, inserted on the 
peduncle and enclosing the inflorescence before anthesis, caducous; staminate 
flower without a perianth, or the perianth stipitate, oblique, with a multidentate 
orifice; stamens numerous, the filaments confluent with the perianth tube and with 
one another, free above, the anthers adnate to the filament, ovoid or linear, de- 
hiscent by longitudinal slits; pistillate perianth none or of 4 short, distinct or 
connate segments, these often confluent with those of adjacent flowers and form- 
ing a persistent multidentate annulus, often accrescent after anthesis and in fruit 
indurate; staminodia in the pistillate flower usually 4, short or long and flexuous; 
ovary free or immersed below in the axis of the spadix and adnate to it, depressed- 
quadrate, truncate or 4-lobate at the apex, 1-celled; style none or short and py- 
ramidal, the stigmas solitary, or 4, depressed, and stellately spreading; ovules 
numerous, the placentae parietal or pendulous from the apex of the cell; fruit a 
fleshy syncarp, composed of distinct or confluent berries crowned by the depressed 
stigmas, the placentae swollen and fleshy; seeds numerous, small, sessile or pendu- 
lous, on a short funicle, the testa fleshy; endosperm fleshy and oily or corneous, 
the embryo minute, straight or curved. 

The family consists of 4 genera of plants confined to tropical 
America. In general appearance they suggest palms, and are likely 
to be confused with that group by one unfamiliar with the trop- 
ical flora. 

Leaves with 2 stout ribs 4-5 mm. broad near the base and extending nearly the 
length of the segments, much more conspicuous than any other nerves, the 
leaf surface mostly smooth and not conspicuously ribbed or nerved ; flowers 
arranged in rings, these alternately staminate and pistillate; perianth none in 
the staminate flower; sap milky Cyclanthus 

Leaves with the 2 or 3 stoutest ribs at most 1.5-2 mm. broad near the base, scarcely 
or not at all more conspicuous than other nerves, the leaf surface with numer- 
ous conspicuous ribs or nerves protruding above the leaf surface; flowers not 
in rings, the staminate in clusters of 4 around a solitary pistillate flower; 
perianth present in the staminate flower; sap watery Carludovica 



300 FIELDIANA: BOTANY, VOLUME 24 

CARLUDOVICA Ruiz & Pavon 

Acaulescent terrestrial herbs, or epiphytic herbs or shrubs, often with greatly 
elongate and scandent stems rooting at the nodes; leaves alternate or fasciculate, 
rarely distichous, bifid or palmately parted, very rarely entire, the segments multi- 
costate, entire or dentate, the short or elongate petiole vaginate-sheathing at the 
base; spadix oblong or cylindric; spathes 2 or more, caducous, concave; flowers 
arranged in dense spirals, 4 staminate ones surrounding a single pistillate flower; 
perianth of the staminate flower stipitate, oblique, fleshy, multidentate; stamens 
numerous, the anthers didymous, oblong, exserted, dorsifixed; pistillate flower free 
or immersed in the rachis and adnate to it, the perianth 4-ahgulate, 4-fid, or 
4-parted, the segments or lobes obtusely quadrate, persistent and indurate; stami- 
nodia 4, greatly elongate, filiform, deciduous; stigmas 4, sessile, stellately spread- 
ing; ovules very numerous. 

The genus consists of perhaps 50 species, widely dispersed in trop- 
ical America. Some are found in Central America, especially in 
Costa Rica and the mountains of Panama, where the species are 
more numerous and the plants more conspicuous and abundant. A 
few species occur in southern Mexico. In Guatemala the number of 
species is small, and the plants are rarely conspicuous. In some re- 
gions of Central America the long and flexible stems of the scandent 
species are used for making baskets and a kind of furniture similar 
to that made from willow (Salix) branches. The ripe fruits of some 
species are intensely fragrant at maturity and juicy. They are eaten, 
at least in the southwestern part of Guatemala, by the Indians. The 
generic name commemorates Carlos XI of Spain and his queen, Luisa. 

Leaves 3-4-parted, the petioles mostly 1-2 meters long. Plants always terrestrial. 

C. palmata 
Leaves bifid, the petioles much shorter. 

Leaf segments mostly 8-15 cm. wide and 40-95 cm. long; fruiting spadix usually 

6-8 cm. long C. utilis 

Leaf segments 1-8 cm. wide, 10-40 cm. long; fruiting spadix 4 cm. long or less. 

Leaf segments mostly 1-3 cm. wide; spathes 6-8 C. microphylla 

Leaf segments mostly 3-8 cm. wide; spathes 2 C. microcephala 

Carludovica microcephala Hook. f. Bot. Mag. pi. 7263. 1892. 

Wet, mixed forest, growing almost always along or near stream 
banks, frequently on limestone, 1300 meters or lower; Alta Verapaz; 
Izabal; Quezaltenango. Honduras; Costa Rica; Panama. 

Plants always terrestrial, small, the caudex very short or practically none, the 
leaves few; petioles slender, elongate, 40 cm. long or shorter, the blades deeply 
bifid, firmly membranaceous, the segments narrowly oblong-lanceolate, 15-40 cm. 
long, mostly 3-8 cm. wide, long-attenuate, 8-24-nerved; peduncles shorter than 
the petioles, slender, with 2 brown lanceolate appressed sheaths at the base; spadix 
small, in fruit 4 cm. long or less, oblong, obtuse, with only about 8 groups of flowers; 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 301 

spathes 2, pale green, ovate to broadly lanceolate, 2.5-6.5 cm. long; staminodia 
3.5-4.5 cm. long. 

Carludovica microphylla Oerst. Vid. Medd. Kjoebenhavn 1857: 
197. 1858. 

Wet, mixed forest, 100 meters or lower; Izabal (base of Sierra del 
Mico). Costa Rica; Panama. 

Usually an epiphytic vine with long, slender stems, ascending tall trees, rarely 
terrestrial along rocky stream banks; leaves short-petiolate, bilobate to below the 
middle, the segments linear-lanceolate, abruptly caudate-acuminate, usually 4-8- 
nerved, mostly 10-15 cm. long; peduncles somewhat shorter than the petioles; 
spadix green, 2.5-3 cm. long. 

Carludovica palmata Ruiz & Pavon, Syst. Fl. Peruv. 291. 1798. 
Cald (Pancajche", Alta Verapaz) ; Palmilla; Palmero; Pojdm (Huehue- 
tenango) ; Jiraca. 

Moist or usually wet, mixed, lowland or mountain forest, some- 
times in open places, especially in second growth, 800 meters or 
lower; Pete*n; Alta Verapaz; Izabal; Suchitepe'quez; Retalhuleu; San 
Marcos; Huehuetenango; often planted for ornament. Southern 
Mexico to Panama; extending southward to Peru. 

Plants terrestrial, acaulescent, often forming large clumps or colonies, the 
slender stiff green petioles 1-2 meters tall, the blades 3-4-parted, a meter broad 
or smaller; inflorescences much shorter than the petioles, pedunculate; spadix 10- 
20 cm. long, red at maturity; staminodia very conspicuous in anthesis, as much 
as 15 cm. long, hairlike, white. 

Called "junco" in Honduras; "soyacal" (Tabasco). The Panama 
hat palm is a well-known and easily recognized plant of the rain for- 
ests of Central America, where the fiber is used in the manufacture 
of hats and other textiles. The fiber, or more properly narrow strips 
of the very young leaves, is notable for its strength, durability, and 
flexibility. From it are made the well-known Panama hats in Cen- 
tral America "sombreros de Jipijapa" which, despite their English 
name, are not made in Panama but in a restricted area of Ecuador 
where atmospheric conditions are particularly suitable for their han- 
dling. These hats are imported in quantity into Guatemala and 
other Central American countries, and are so esteemed that poor 
people often buy them at very high prices. Because of its handsome 
appearance, the Panama hat palm is grown commonly for orna- 
ment in parts of Guatemala where it is not native, and at rather 
high elevations of 1500 meters or more, far above the zones where 
it grows wild. 



302 FIELDIANA: BOTANY, VOLUME 24 

Carludovica utilis (Oerst.) Benth. & Hook, ex Hemsl. Biol. 
Centr. Amer. Bot. 3: 416. 1885. Sarcinanthus utilis Oerst. Vid. 
Medd. Kjoebenhavn 1857: 197. 1858. Tepejilote (Quezaltenango; 
probably an erroneous name); Cuajiote (Huehuetenango). Figure 53. 

Wet, mixed forest of the mountains or lowlands, mostly at 300- 
1500 meters; Alta Verapaz; Izabal; Solola; Suchitepe"quez; Quezalte- 
nango; San Marcos; Huehuetenango. British Honduras to Panama. 

Plants terrestrial and acaulescent or often epiphytic or scandent and with 
much elongate stems; petioles slender, equaling or shorter than the blades; leaf 
blades bifid to below the middle, mostly 40-95 cm. long, with 3 main costae, the 
segments 7-15 cm. wide or wider, abruptly short-acuminate, 8-32-nerved; pedun- 
cles much shorter than the petioles; spathes 5, remote, 5-10 cm. long; spadix 
oblong, with numerous groups of flowers, rounded at the apex. 

The leaves were used formerly for weaving hats similar to Pan- 
ama hats, and probably they still are used locally in this manner. 
The stems of the epiphytic plants are stout, very flexible, and tough. 



GYGLANTHUS Poiteau 

Plants terrestrial, acaulescent, with milky sap; leaves arranged spirally, petio- 
late, the blades deeply bifid; spadix subtended by 5-7 petaloid or somewhat folia- 
ceous, large spathes; staminate and pistillate flowers borne on the same spadix, in 
separate alternating whorls, the flowers of each whorl adnate and confluent; stami- 
nate flowers naked; pistillate flowers enclosed by 2 rings of fleshy tissue representing 
the concrescent perianths; staminodia numerous, short; fruit a syncarp, consisting 
of the several fleshy whorls of pistillate flowers, the whole suggestive of a large screw. 

The genus consists of two or three species, all tropical American. 
Only the following is known from North America. 

Cyclanthus bipartitus Poit. Me"m. Mus. Paris 9: 35. pi. 2, 3. 
1822. Cola de pava; Palmiche. 

Wet, mixed, lowland forest, 350 meters or lower; Alta Verapaz; 
Izabal; Huehuetenango. Honduras; Costa Rica; Panama; south- 
ward to Peru and Brazil. 

Plants cespitose, often forming large dense clumps, 1-2 meters high; petioles 
50-100 cm. long, the blades of the same length or longer, bifid almost to the base, 
thick, not plaited or conspicuously costate, the segments broadly linear-lanceolate, 



FIG. 53. Carludovica utilis. a, Staminodia (X 2). b, Stamen (X 15). c, 
Portion of flowering shoot with leaf (X J4). d, Surface of pistillate flower, from 
above (X 3^)- e, Staminate flower, lateral view (X 3%). /, Inflorescence, 
showing spathes (X %) 




303 



304 FIELDIANA: BOTANY, VOLUME 24 

7-15 cm. wide, acute or acuminate, with a stout costa; peduncles elongate, arising 
from the base of a cluster of leaves; spathes 4-5, 20-22 cm. long, green, longer than 
the spadix, the inner ones greenish white, oblong-lanceolate, cuspidate; spadix with 
8-12 whorls of pistillate flowers, in fruit as much as 22 cm. long and 5 cm. thick. 



ARACEAE. Arum Family 
Reference: Engler & Krause, Pflanzenreich IV. 23. 

Plants terrestrial or more often epiphytic, usually glabrous, acaulescent or with 
erect or scandent stems, often with tuberous roots; leaves petiolate, entire, lobate, 
or parted, the petioles sheathing at the base; reduced leaves or cataphylls often 
present on the stems; peduncles simple, axillary or terminal; flowers small, perfect 
or unisexual, arranged on a terete elongate spadix, the pistillate flowers below, the 
staminate above, the spadix subtended by a foliaceous spathe, this persistent or 
deciduous in fruit; perianth none, or of 4-6 segments, these distinct or connate; 
stamens 1-many, distinct, hypogynous; ovary usually entire, 1-many-celled; style 
none or present and elongate; ovules 1 or more in each cell; fruit baccate, the berries 
free or often connate, containing 1-many seeds. 

More than 100 genera, widely dispersed, chiefly in the tropics. 

The Araceae or aroids are one of the largest families of mono- 
cotyledonous plants, the species being distributed most abundantly 
in tropical regions, with a few species extending into the temperate 
belts. Most of the American ones are epiphytes, some of them acau- 
lescent plants, others large vines that often form mantles of showy 
leaves over the trunks or branches of trees. Their leaves are usually 
thick and fleshy, and the plants are able to withstand dryness better 
than many other epiphytes, yet they seldom are found in dry regions. 
The leaves of many aroids have attractive form and texture, and 
when well developed many of the species are highly ornamental. 
Some of them, such as the Monsteras and Philodendrons, form a con- 
spicuous element of the epiphytic vegetation characteristic of trop- 
ical regions. 

In some genera, especially those consisting of scandent plants, 
there is a remarkable difference in leaf form between the juvenile 
and the adult stages. This variation is especially marked in such 
groups as Philodendron, Monstera, and Syngonium. Juvenile plants 
often are so different from adult ones that it is difficult to associate 
the various forms. A careful study of the life histories of the Cen- 
tral American scandent Araceae would give interesting results, and 
perhaps make the determination of the juvenile forms less difficult. 

Many or most of the aroids contain in their tissues needle-like 
crystals of calcium oxalate (at least it is usually so designated). 
When any portion of the foliage is chewed, these crystals penetrate 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 305 

the mouth tissues, often causing intense irritation and swelling. In 
the case of the Araceae used for food such as Xanthosoma and Mon- 
ster a special care must be taken in their preparation. Fruiting spa- 
dices (such as those of Monstera) must be fully ripened, and tubers 
or rootstocks must be thoroughly exposed to heat in order to elimi- 
nate the pernicious crystals. 

Many of the Guatemalan plants of this family, especially Mon- 
stera and some species of Philodendron, produce long, flexible, very 
tough aerial roots that attain a great length when the plants grow 
high on trees. These roots are used very commonly in Guatemala 
and other parts of Central America for making the so-called mimbre 
furniture. While still moist the roots are wrapped tightly about 
frames of chairs and other articles of furniture, and as they dry they 
become still tighter, thus resembling similar furniture made in the 
United States from other material. The industry is locally an im- 
portant one. These roots sometimes reach the United States, and 
we have seen samples of them from the Chicago customs house. 

Plants aquatic, floating on the surface of water; leaves arranged in a rosette, 

spongy Pistia 

Plants not aquatic; leaves not forming a rosette, not spongy. 
Leaves peltate. Plants with tuberous rhizomes, cultivated or escaped, always 
terrestrial. 

Leaves green, not spotted Colocasia 

Leaves green but spotted with white, cream, pink or other colors . . .Caladium 
Leaves not peltate. 

Leaf blades perforated, sometimes also pinnatifid Monstera 

Leaf blades never perforated. 

Leaves digitately compound, pedately 3-11-cleft, or pinnatifid. 

Blades pinnatifid Philodendron 

Blades trisect or pedately or digitately 5-11-parted. 

Flowers all perfect, the spadix uniform; perianth present. Leaves 
digitately 5-11-parted. (One Guatemalan species.). . .Anthurium 
Flowers unisexual, the lower part of the spadix with pistillate flowers, 
the upper with staminate flowers; perianth none. 

Plants terrestrial, acaulescent Xanthosoma 

Plants epiphytic, with elongate caudices. 

Spadix almost equaling the spathe; leaf blades 3-parted. (Two 
Guatemalan species.) Philodendron 

Spadix much shorter than the spathe; leaves trisect or pedately 
5-11-cleft Syngonium 

Leaves entire, but often hastate or cordate and with basal lobes. 
Flowers all or mostly perfect, the spadix uniform, not divided into a 

pistillate and staminate portion. 
Plants terrestrial, acaulescent. 



306 FIELDIANA: BOTANY, VOLUME 24 

Leaves acute to rounded at the base Spathiphyllum 

Leaves hastate, with elongate basal lobes Urospatha 

Plants normally epiphytic, often with elongate caudices. Spathe 
spreading from the base of the spadix, not enclosing it, usually 
deciduous. 

Stigma discoid; plants scandent or acaulescent Anthurium 

Stigma oblong or linear; plants more or less scandent, the caudices 

with elongate internodes. 
Peduncle cernuous or recurved at the apex at or before anthesis. 

Stenospermation 
Peduncle erect at the apex, never cernuous or recurved. 

Peduncles shorter than the spat he Monster a 

Peduncles much longer than the spathe Rhodospatha 

Flowers unisexual, the spadix divided into a pistillate and staminate por- 
tion, these contiguous or separated by a sterile portion. Flowers 
without a perianth. 
Plants epiphytic, at least normally so, with usually scandent stems, 

these rooting at the nodes Philodendron 

Plants terrestrial, acaulescent or with erect stems supported by prop 

roots. 

Leaf blades acute to rounded or subcordate at the base . Dieffenbachia 
Leaf blades deeply cordate or hastate at the base. 

Peduncles equaling or exceeding the leaves. Plants acaulescent, 

cultivated or rarely naturalized Zantedeschia 

Peduncles much shorter than the leaves. 

Stamens distinct; plants with erect caudices, these usually sup- 
ported by prop roots, often or usually prickly. 

Montrichardia 

Stamens connate or coherent; plants acaulescent or with pros- 
trate caudices erect at the apex, without prop roots, 
unarmed . . . . Xanthosoma 



ANTHURIUM Schott 

Reference: Engler, Pflanzenreich IV. 23B: 53. 1905. 

Plants usually epiphytic, the caudex usually very short and the plants appear- 
ing almost or quite acaulescent, the stems rarely elongate, the internodes short or 
elongate; petiole short or elongate, short- vagina te at the base, terete or more or less 
canaliculate, geniculate near the apex and more or less swollen at the node, the 
blades commonly thick, often very thick and coriaceous when dried, variable in 
form, simple or parted or digitate; peduncles usually elongate; spathe persistent, 
usually green or purplish and lanceolate or linear-lanceolate, not enclosing the 
spadix and during anthesis spreading from its base; spadix sessile or stipitate, 
cylindric or conoid or caudiform, densely many-flowered, flowering from the base 
upward, in fruit much thickened; flowers perfect, with a perianth; sepals 4, forni- 
cate at the apex, connivent, sub truncate, accrescent in fruit; stamens 4, the fila- 
ments subcompressed, slightly narrowed above into the connective, equaling the 
sepals; anthers short, the cells ovate or oblong-ovate, extrorse, opening by a longi- 
tudinal slit; ovary ovoid or oblong or obovoid, truncate at the apex or attenuate 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 307 

into the style, 2-celled; ovules 2 or more or often 1 in each cell; style short or none, 
the stigma small, discoid; berries very juicy, variable in shape and color, the cells 
usually 1-seeded; seeds oblong, more or less attenuate to the apex. 

The largest genus of the family, with about 500 species, generally 
distributed in the American tropics. About 65 species are recorded 
for Central America, the majority in Costa Rica and Panama. Most 
of the species grow upon trees, but some upon rocks or even upon the 
ground. They are chiefly small and not very conspicuous plants, but 
some are rather ornamental. 

Leaves digitately compound A. aemulum 

Leaves entire. 

Leaf blades acute to long-attenuate at the base. 

Cells of the ovary normally 2-ovulate; stems usually elongate and more or 
less scandent; leaves small, commonly less than 5 cm. wide, and short. 

A. scandens 
Cells of the ovary 1-ovulate; stems short or obsolete, very rarely elongate; 

leaves small or large. 
Leaves very large, commonly 12-30 cm. wide or broader, not 3-nerved. 

Node of the petiole 3-carinate dorsally A. crassinervium 

Node of the petiole not carinate dorsally, flat. 

Upper joint of the petiole, above the node, cubical A. tetragonum 

Upper joint of the petiole longer than broad A. Schlechtendalii 

Leaves relatively small, mostly less than 8 cm. wide and often 3 cm. or 

narrower, often conspicuously 3-nerved. 

Leaf blades evidently broadest above the middle, long-attenuate to the 
base; axis and upper part of peduncle of spadix slender, mostly 

11.5 mm. thick A. scolopendrinum 

Leaf blades broadest at or below the middle, not long-attenuate to the 
base; axis and upper part of peduncle of spadix broader, 2-3 mm. 
thick. 

Plants acaulescent or nearly so; leaves mostly 20-35 cm. long or longer. 

A. Bakeri 
Plants with much elongate stems; leaves mostly 11-15 cm. long. 

A. Pittieri 

Leaf blades rounded to deeply cordate at the base. 

Leaf blades deeply cordate at the base, with large and conspicuous basal lobes. 
Cells of the ovary normally 2-ovulate; spathe bright red . . A. Scherzerianum 
Cells of the ovary 1-ovulate; spathe not red, usually green, white, or pur- 
plish, or suffused with rose. 
Spadix 12-20 cm. long or longer. 

Spathe linear-lanceolate, 0.4-1.2 cm. wide; spadix about 8 mm. thick 

or narrower; leaves mostly 12-22 cm. wide A. verapazense 

Spathe lanceolate, 2-6.5 cm. wide; spadix 8-20 mm. thick; leaves 

mostly 25-60 cm. wide. 
Spadix 3-4.5 cm. stipitate; peduncles 2 cm. thick near base. 

A. titanium 
Spadix subsessile, mostly 0.3-1 cm. stipitate; peduncles mostly 

0.3-1 cm. thick near base A. concinnatum 

Spadix usually much less than 10 cm. long. 



308 FIELDIANA: BOTANY, VOLUME 24 

Basal lobes of the leaves separated by a very broad and open sinus; 

usually at 1800-2700 meters altitude A. montanum 

Basal lobes of the leaves separated by a narrow oblong sinus, or the 
sinus closed by the somewhat overlapping lobes. 

Spathe 1 cm. wide or narrower A. seamayense 

Spathe 3-5 cm. wide. 

Leaf blades subcoriaceous; spathes mostly white, not evidently 
nerved, opaque and thickish when dried, oblong, not con- 
spicuously amplexicaul; at 1000-1800 meters altitude. 

A. fraternum 

Leaf blades firmly membranaceous; spathes pale green, conspicu- 
ously nerved, thin when dried, ovate to lance-ovate, conspicu- 
ously amplexicaul A. silvigaudens 

Leaf blades rounded to truncate or rounded and shallowly cordate or emargi- 
nate at the base, never with large basal lobes. 

Leaves, when dried, very thick and coriaceous, the costa and primary nerves 

very stout and prominent. 
Leaf blades subcordate at the base, with small but evident basal lobes, the 

blade gradually narrowed from the base to the apex A. Seleri 

Leaf blades rounded or truncate at the base, without basal lobes. 

Leaf blades, including the costa, conspicuously 5-7-nerved at the base 
on the lower surface, the lowest 2-3 pairs of lateral nerves con- 
spicuous; leaf blades broadest at the base or lower half; spadix 
conspicuously stipitate A. subcordatum 

Leaf blades with only the prominent midrib showing on the lower sur- 
face, the lateral nerves similar throughout; leaf blades of about the 
same width throughout or broadest at the middle; spadix sessile. 

Leaves green beneath, less than 3 times as long as wide, 12-18 cm. 
wide, conspicuously reticulate-nerved on both sides, the nerves 
prominently elevated below and sulcate above A. retiferum 

Leaves glaucous or glaucescent beneath, more than 4 times as long 
as wide, 6-14 cm. wide, not conspicuously reticulate-nerved, the 
nerves not prominently elevated below or sulcate above. 

A. parvispathum 

Leaves, when dried, thin and almost membranaceous, the costa and primary 
nerves very slender. 

Plants acaulescent or nearly so A. subcordatum 

Plants with elongate, creeping or scandent stems. 

Leaf blades rather deeply cordate at the rounded base; spadix con- 
spicuously stipitate A. myosuroides 

Leaf blades broadly rounded or broadly subcordate at the base; spadix 
subsessile A. Tuerckheimii 

Anthurium aemulum Schott, Bonplandia 165. 1859. Okil 
(Pet&i, Maya?). 

Moist or wet, lowland forest, 750 meters or lower; Pete*n; Alta 
Verapaz; Izabal; Santa Rosa; Escuintla; Retalhuleu. Southern 
Mexico; British Honduras to Panama. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 309 

An epiphytic vine, often very large, the thick caudex with long internodes; 
petioles long and slender; leaf blades digitately 5-11-parted, the segments sessile 
or petiolulate, oblong-elliptic to oblanceolate, 20-30 cm. long in adult leaves, cau- 
date-acuminate, narrowed to the base, thin when dried; peduncles stout, 5-10 cm. 
long; spathe lanceolate, green, 10 cm. long, 2.5 cm. wide at the base; spadix dull 
green, short-stipitate, about 5 cm. long and almost 2 cm. thick. 

A conspicuous and rather handsome plant, very different from 
other Guatemalan species in its digitately parted leaves. 

Anthurium Bakeri Hook. f. Bot. Mag. pi. 6261. 1879. 

Usually epiphytic in dense wet forest, 1500 meters or lower, 
chiefly at very low elevations; Pet&i; Alta Verapaz; Izabal; Hue- 
huetenango. British Honduras; Nicaragua; Costa Rica. 

Plants acaulescent or sometimes developing a short, somewhat woody caudex, 
the caudex with very short internodes; petioles slender or stout, mostly one-fourth 
as long as the blades or shorter, rarely more elongate, dilated at the base; leaf 
blades narrowly elliptic-lanceolate, 30-50 cm. long, usually 3-6 cm. wide, thick and 
somewhat coriaceous when dried, acuminate, acute or attenuate at the base, widest 
at or near the middle, green above, paler green beneath, conspicuously 3-nerved, 
the lateral nerves very numerous and prominent, especially beneath; peduncles 
mostly slender and elongate, half as long as the leaves or sometimes equaling them; 
spathe green or bronze, usually 3.5-4.5 cm. long and 1-1.5 cm. wide, often smaller, 
usually rounded and apiculate at the apex, reflexed, decurrent at the base; spadix 
short-stipitate or subsessile, commonly 4-5 cm. long but sometimes 9 cm. long in 
an thesis, greenish, whitish, or purplish, very slender in anthesis, in fruit often more 
elongate (in the type as much as 20 cm. long and 3 cm. thick in the fresh state) ; 
berries bright red at maturity, ovoid, as much as 1 cm. long in the fresh state but 
much smaller when dried. 

The local material is somewhat variable, and ampler collections 
may necessitate its division into two or more taxa. The alleged dif- 
ferences between A. Bakeri and A. consobrinum Schott (1855) do not 
appear to be stable ones, leaving considerable doubt whether they 
can be maintained as distinct species. They vary greatly in shape of 
leaf blade, and in measurements of spathe and spadix. If eventually 
treated as one species, the name A. consobrinum has priority. 

The Guatemalan material was referred first to A. Friedrichsthalii 
Schott, which was based on material collected by Friedrichsthal at 
"Cativo," said to be in Guatemala. No such locality is known in 
Guatemala, and it seems probable that the collection actually was 
made in the Panama Canal Zone. 

Anthurium concinnatum Schott, Prodr. 522. 1860. A. bogo- 
tense Schott var. concinnatum (Schott) Engler, in DC. Mon. Phan. 
2: 184. 1879. 



310 FIELDIANA: BOTANY, VOLUME 24 

Epiphytic or terrestrial in dense, wet forest, generally 75-1100 
meters, ascending to 2000 meters in the Sierra de las Minas; Izabal; 
Zacapa; Chiquimula; Alta Verapaz; Huehuetenango. British Hon- 
duras; Honduras; Costa Rica; Panama. 

Plants usually large and coarse; petioles 3.5-9 dm. long, geniculate 2-3 cm. 
below the base of the blade, the node 3-7 mm. broad; leaf blades subcoriaceous, rich 
green and shining above, pale green below, broadly ovate-cordate or triangular- 
cordate, 2.5-7 dm. long, 13-43 cm. wide, obtuse to rounded at the apex and 
abruptly long cuspidate-acuminate, deeply cordate at the base with a very broad 
and open sinus, entire, 7-nerved from the base, the lowest 2 nerves emitting usually 
5 prominent nerves on the lower surface of the broadly rounded postical lobes, the 
primary lateral nerves slender, forming an irregular collective nerve 4-10 mm. from 
the margins, the veins prominulous, laxly reticulate; peduncle up to 5 dm. or more 
long, 0.3-1 cm. or more thick near the base; spathe coriaceous or subcoriaceous, 
spreading to reflexed, green, greenish-white or whitish, broadly lanceolate, abruptly 
cuspidate-acuminate, 6-25 cm. long, 2-5.5 cm. wide near the base; spadix in an the- 
sis pale or dull green, lilac-buff, pink, whitish, or brownish, in fruit reddish, 6.5-24 
cm. long, 8-18 mm. thick near the base, of about the same diameter throughout, 
slightly narrower above, usually longer than the spathe, subsessile or on a stipe 
usually 3-10 mm. long, rarely longer, and 3-7 mm. thick; fruits oblong or oblong- 
ovoid. 

Anthurium crassinervium (Jacq.) Schott, Melet. 1 : 22. 1832. 
Pothos crassinervia Jacq. Icon. 3: pi. 609. 1793. A. Salviniae Hemsl. 
Diag. PL Mex. 36. 1879; Biol. Centr. Am. Bot. pi. 99. 1888. Que- 
quesquillo; Colmenero; Baatun, Ucutzh box (Pete*n, Maya) . Figure 54. 

On trees in wet forest, sometimes on wet rocks or rarely (prob- 
ably by accident) terrestrial, 1400 meters or lower, most common at 
low elevations; Pete"n; Alta Verapaz; Izabal; Jalapa; Santa Rosa; 
Escuintla; Sacatepe"quez; Retalhuleu; San Marcos. Southern Mex- 
ico; British Honduras to Panama; northwestern South America. 

Plants very large, usually epiphytic, acaulescent or nearly so, the caudex thick; 
petioles very thick and stout, mostly 15-30 cm. long and 2-3 cm. in diameter, 
geniculate 1.5-2 cm. below the base of the blade, the joint as broad as long or 
broader; leaf blades somewhat coriaceous when dried, thick and succulent when 
alive, oblong-lanceolate or oblong-obovate, commonly 60-100 cm. long and 25-30 
cm. wide, or sometimes even larger, acute and cuspidate, gradually narrowed to 
the base, the margins often undulate, the costa very thick and prominent, the pri- 
mary nerves 9-14 on each side; peduncles 50-70 cm. long, about 1 cm. thick, 
pendent or recurved in fruit; spathe thick, bronze-green or bronze-purple, broadly 
linear-lanceolate, long-acuminate, clasping at the base, 6-30 cm. long and 1.5-3.5 
cm. wide; spadix short-stipitate or sessile, caudiform, purplish-green, 4-30 cm. 
long, 0.5-1.5 cm. thick, in fruit becoming 4-5 cm. or more in diameter; berries 
ovoid, scarlet, 1 cm. long; seeds 4 mm. long, minutely verruculose. 

Sometimes called "hoja de piedra" in Honduras. The plant is 
perhaps the showiest of all local species of Anthurium because of its 




FIG. 54. Anthurium crassinervium. a, Portion of surface of spadix (X ' 
b, Pistil (X 9). c, Stamen (X 9). d, Longitudinal section through ovary (X 9). 
e, Sepal (X 9). /, Habit (X Ve). g, Flower with spathe and spadix (X ^). 

311 



312 FIELDIANA: BOTANY, VOLUME 24 

huge leaves, which usually are numerous and form a large cluster. 
The inflorescences are not conspicuous, but the very large and heavy 
fruiting spikes that hang from the base of the plant attract attention 
because of their vivid coloring. The plant does well in cultivation 
and thrives even when planted in the ground, as we observed it at 
Quirigua. 

There is considerable variation in the width, shape, and base of 
the leaf blade. Brenes 12639 from Costa Rica shows a rounded tend- 
ency at the base of the leaf blade. In many plants the lower part of 
the leaf blade is narrowly attenuate to the petiole. In other collec- 
tions, as in Standley 76313 from the vicinity of Jutiapa, Guatemala, 
Standley 58294- from near Las Lajas, Dept. Sacatepe"quez, Guate- 
mala, Standley 88391 from Dept. Retalhuleu, Guatemala, and Stand- 
ley 76313 from the vicinity of Jalapa, Guatemala, the leaf blade is 
relatively broad in the middle and tapers to a somewhat rounded 
base, best developed in Standley 76313. Probably A. Salviniae was 
described from a plant of the latter type, and from such collections as 
Elias 1404 and 1477 from Colombia, with an extreme development of 
the rounded base. Actually, Standley 58294, from near Las Lajas at 
1200 meters elevation, is not too far from the type locality of A. Sal- 
viniae, collected on Volcan de Fuego at 1200 meters either in the 
department of Sacatepe"quez or Chimaltenango. Engler cites a col- 
lection of Heyde & Lux 4278 from Naranjo, Dept. Santa Rosa, taken 
at 1100 meters, and Standley 78060, taken near Cuilapilla of the same 
department at an elevation of 900 meters, has leaves with slightly 
rounded bases, which begin to approach those of the type. Other 
collections of A. crassinervium were obtained by Standley from the 
lower slopes of the Volcan de Fuego, the type locality of A. Salviniae. 
When the A. crassinervium complex has been studied with greater 
detail, it may be that A. Salviniae will prove distinct. 

Anthurium fraternum Schott, Oesterr. Bot. Wochenbl. 293. 
1857; Hemsl. Biol. Centr. Amer. Bot. pi. 97. Gusnay; Gusnay de 
monte; Huisnay. 

Moist or wet forests of the Pacific boca costa, 1000-1800 meters; 
endemic; Escuintla; Sacatepe"quez (type from Barranco Hondo, base 
of Volcan de Fuego, Salvin); Chimaltenango; Solola; Suchitepe"quez ; 
Quezaltenango; San Marcos; to be expected in Chiapas. 

Plants usually epiphytic, sometimes on rocks, acaulescent or nearly so, the 
caudex almost 1 cm. thick; petioles short- vaginate, equaling or longer than the 
blade; leaf blades elongate-ovate-cordate, mostly 20-40 cm. long and 12-21 cm. 
wide, acuminate, deeply cordate at the base, the sinus relatively narrow, the basal 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 313 

lobes rounded and somewhat incurved, the primary costal nerves about 5-6 on 
each side, prominent and rather stout, united to form a slender submarginal nerve; 
peduncles 20-30 cm. long or more; spathe subcoriaceous, subopaque, oblong or sub- 
elliptic, 4.5-10 cm. long and 1.5-3.5 cm. wide, caudate-acuminate, usually white or 
becoming green, clasping at the base; spadix borne on a stipe 5-15 mm. long, 5-8 
cm. long and 8 mm. thick, or in fruit as much as 10 cm. long and 2 cm. thick, green 
or purplish. 

Anthurium montanum Hemsl. Diag. PL Mex. 36. 1879; Biol. 
Centr. Amer. Bot. pi. 98. Cartucho. 

Common in many regions in moist or wet forest, usually at 1800- 
2900 meters; endemic; Chiquimula; Sacatepe"quez; Chimaltenango 
(type collected on Volcan de Fuego above Calderas, Salviri); Solola; 
Quiche"; Huehuetenango; Suchitepe"quez; Quezaltenango; San Marcos. 

Plants acaulescent or nearly so, usually epiphytic, the caudex 1.5 cm. thick, 
with very short internodes; petioles slender, equaling or longer than the blades; 
leaf blades subcoriaceous, elongate-cordate, ovate-cordate, or triangular-cordate, 
22-35 cm. long, 9-23 cm. wide, acuminate, deeply cordate at the base, the sinus 
very broad and open, the basal lobes large and rounded, directed downward or 
slightly outward; peduncles slender, 20-30 cm. long; spathe oblong, pale green or 
reddish green, 4.5-8.5 cm. long, (1-) 1.5-3 cm. wide, long-acuminate or cuspidate; 
spadix short-stipitate, 2.5-7 cm. long, 5-8 mm. thick, very obtuse, dark purple. 

It is somewhat doubtful whether this species is really distinct from 
A. fraternum. Both are frequent in the same general region, but 
A. fraternum usually occurs at the lower elevations on the slopes of 
the volcanoes, from 1000-1800 meters, while A. montanum is gener- 
ally encountered at higher altitudes of 1800-2700 meters. They may 
be variants of a single species ranging from the lower to the higher 
elevations, or may possibly cross with each other at intermediate alti- 
tudes. They are favorite pot plants in gardens and patios of the cen- 
tral uplands and like most epiphytic aroids, they thrive when potted 
in ordinary soil. They ornament many patios, withstand neglect, 
resist drouth, and even do well for a long time inside the house in 
rooms that receive no sunlight. 

Anthurium myosuroides (HBK.) Endl. Gen. 240. 1837. Pothos 
myosuroides HBK. Nov. Gen. & Sp. 1: 62. pi. 18. 1815. Corazonado; 
Contin; Chile de mico (fide Aguilar). 

Moist or wet forest, chiefly on the Pacific slopes of the volcanoes, 
1000-1400 meters; Chimaltenango; Quiche"; Quezaltenango; San Mar- 
cos. Southern Mexico(?); British Honduras to Panama; Colombia. 

Plants epiphytic, the caudex very slender and usually much elongate, the in- 
ternodes often 5-7 cm. long; petioles slender, mostly 10-15 cm. long, the blades 



314 FIELDIANA: BOTANY, VOLUME 24 

ovate-elliptic or ovate, 12-15 cm. long, 7-8 cm. wide, gradually acuminate or cus- 
pidate-acuminate, rounded at the base and shallowly cordate, the sinus open or 
almost closed, the primary costal nerves 9-10 on each side, oblique, united to form 
a conspicuous collective nerve remote from the margin; peduncles slender, 6-15 cm. 
long; spathe lanceolate or linear-lanceolate, 5-10 cm. long, green, long-attenuate; 
spadix green, caudiform, short-stipitate, 5-12 cm. long, about 4 mm. thick in an- 
thesis; berries bright red, almost 1 cm. long. 

Called "Tietie" in British Honduras. 

Anthurium parvispathum Hemsl. Biol. Centr. Amer. Bot. 3: 
432. 1885. A. hypoleucum Standl. Journ. Wash. Acad. Sci. 17: 246. 
1927 (type from Santa Rosa, Baja Verapaz, Tuerckheim 11.2214). 
A. brevipetiolatum Engler ex Krause, Notizbl. Bot. Gart. Berlin 11: 
610. 1942 (based on the same collection as A. hypoleucum). Madre 
de arpdn (fide Aguilar). 

Usually on moist or dry, shaded or exposed rocks, about 1500 
meters; endemic; Baja Verapaz (type from Santa Rosa, Salvin & 
Godman 408); Quiche*. 

Plants acaulescent or nearly so, the caudex short and very thick; petioles two- 
thirds as long as the blades, or sometimes longer, very stout, geniculate just below 
the base of the blade; leaf blades very thick and fleshy when alive, thick-coriaceous 
when dried, oblong-lanceolate, 30-60 cm. long, 6-14 cm. wide, usually obtuse and 
apiculate at the apex but sometimes acute or acuminate, obtuse to subtruncate at 
the base, deep green above, pale or sometimes whitish beneath, the primary costal 
nerves 8-9 on each side, remote, united near the margin to form an evident collec- 
tive nerve, the costa thick and stout, very prominent beneath; peduncles as much 
as 60 cm. long; spathe oblong or ovate-oblong, usually much shorter than the 
spadix, 2.5-7 cm. long, 8-12 mm. wide, green or bronze; spadix sessile, as much as 
9 cm. long, 6-10 mm. thick in anthesis, in fruit 2 cm. or slightly more in diameter, 
bronze. 

This striking and ornamental plant is abundant and conspicuous 
on large rocks along the road in the Santa Rosa region, where it 
often forms large clumps or colonies. It is notable because it grows 
in full sunlight, which may be intense and hot in this locality, rather 
than in forest shade, and its very thick and hard leaves, exceedingly 
resistant to desiccation, fit it well for such a habitat. Its range, as 
now known, is very limited for a plant of this family. 

Anthurium Pittieri Engler, Bot. Jahrb. 25: 373. 1898. A. radi- 
cosum Standl. & Steyerm. Field Mus. Bot. 23: 210. 1947. 

Epiphytic in wet forest, 300-350 meters; Alta Verapaz (type 
of A. radicosum from Rio Icvolay, north and northwest of Finca 
Cubilgiiitz to Quebrada Diablo, Steyermark 44776). British Hon- 
duras (Pueblo Viejo); Costa Rica; Panama. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 315 

Plants epiphytic, the caudex slender, 3-10 mm. thick, the internodes 7-12 cm. 
long, or the terminal ones shorter; cataphylls caducous, thin, brown, obtuse, about 
4-5 cm. long and 0.5-1 cm. wide; petioles very slender, 4-7 cm. long, short-vagi- 
nate; leaf blades coriaceous alive, very thin and almost membranaceous when dried, 
narrowly elliptic-oblong to oblong-lanceolate, 11-16 cm. long, 2.5-6 cm. wide, 
rather abruptly acuminate, acute or short-acuminate at the base, fuscescent when 
dried, slightly paler and brownish beneath, the primary lateral nerves 7-12 on 
each side, very slender and barely prominulous, inconspicuous, united to form an 
inconspicuous collective nerve close to the margin; peduncles very slender, 10-15 
cm. long; spathe reflexed, pale green, oblong-ovate to linear-lanceolate, 1-2 (up 
to 7) cm. long, 7-10 mm. wide, obtuse and apiculate to acuminate, clasping at the 
base; spadix sessile or nearly so or stipitate, slender, in anthesis 4.5-5 cm. long (in 
fruit up to 8 cm. long), 3-4 mm. thick, not narrowed upward, the flowers rather few. 

Because of the great amount of variation in the dimensions of 
spathe and length of the stipe of the spadix in A. Pittieri, it has been 
found impossible to separate the mass of this variation from A. 
radicosum. 

Anthurium retiferum Standl. & Steyerm. Field Mus. Bot. 23: 
211. 1947. 

Epiphytic or terrestrial in moist Liquidambar forest, 1500-2600 
meters; endemic; Huehuetenango (Sierra de los Cuchumatanes; 
type from Cerro Huitz, between Mimanhuitz and Yulhuitz, Steyer- 
mark 48621; also on Cerro Negro, 2 miles east of Las Palmas). 

Petioles 26-37 cm. long or longer, stout, 6-8 mm. thick, geniculate, 1-1.5 cm. 
below the base of the blade, the node almost 1 cm. thick; leaf blades subcoriaceous 
and rigid when dried, somewhat brownish or fulvescent, oblong-elliptic or narrowly 
oblong-elliptic, 32-42 cm. long, 12-19 cm. wide, acute or subobtuse and apiculate 
at the apex, rounded or very obtuse at the base, somewhat lustrous, concolorous 
or nearly so, the primary lateral nerves about 12 on each surface, impressed on the 
upper surface, slender and salient beneath, the secondary nerves and veins very 
prominent and conspicuously and laxly reticulate; peduncle 19 cm. long, stout, 
4 mm. thick; spathe (a perfect one not seen) reflexed, purple-rose, more than 6.5 
cm. long, about 13 mm. wide, cuspidate-acuminate; spadix sessile, cylindric, 9 cm. 
long, 5 mm. thick, dull purple. 

Anthurium scandens (Aubl.) Engler in Mart. Fl. Bras. 3, pt. 2: 
78. 1878. Dracontium scandens Aubl. PL Guian. 836. 1775. A. 
rigidulum Schott, Oesterr. Bot. Zeitschr. 8: 180. 1858. Elote; Huis- 
nay de montana; Mazorquillo; Cux (Coban, Quecchi). 

Mostly epiphytic in moist or wet, dense forest, 1800 meters or 
lower; Alta Verapaz; Izabal; Zacapa; Chiquimula; Jalapa; Es- 
cuintla; Sacatepe'quez; Chimaltenango; Quiche"; Suchitepe"quez; 
Quezaltenango; San Marcos; Huehuetenango. Southern Mexico; 
British Honduras to Panama; South America. 



316 FIELDIANA: BOTANY, VOLUME 24 

Plants usually epiphytic, sometimes growing on rocks, rarely, probably as a 
result of accident, on the ground, the caudex often greatly elongate and scandent 
or pendent, usually covered by fibrous remains of the cataphylls, the internodes 
mostly elongate; petioles one-fifth to one-half as long as the blades, narrowly cana- 
liculate; leaf blades rather thin when dried, lance-elliptic, mostly 8-13 cm. long 
and 2.5-4.5 cm. wide, acute at each end, or rarely obtuse at the apex, the primary 
costal nerves ascending at an angle of about 45 degrees, united near the margin to 
form a slender and not very conspicuous marginal nerve; peduncles slender, equal- 
ing or as much as 4 times as long as the petioles; spathe green, lanceolate or oblong- 
lanceolate, cuspidulate, somewhat shorter than the spadix, reflexed; spadix pale 
green, slender, short-stipitate, commonly 2-3 cm. long; berries lavender. 

Known in Costa Rica by the names "elotillo," "elotico," and 
"maicillo." Perhaps the commonest and most widely dispersed of 
all Central American species of Anthurium, often cultivated in gar- 
dens or patios. Although normally epiphytic, the plant sometimes 
grows upon logs or on the ground, where probably it persists after 
falling from a tree. The stems seldom are closely attached to the 
trees but more often are free for the greater part of their length, dan- 
gling from the trunks or branches. Illustrated, Pflanzenreich IV. 
23B: /. 21. Engler refers Guatemalan material to two varieties, 
A. scandens var. dolosum (Schott) Engler and A. scandens var. vio- 
laceum (Swartz) Engler, which appear to be varieties of no systematic 
significance. Some Central American material of this group has been 
referred to A. trinerve Miq., which apparently does not differ essen- 
tially from A. scandens. Some of the Guatemalan collections could 
be referred to A. rigidulum Schott. This is maintained by Engler as 
a distinct species but with the curious statement that he does not 
consider it distinct from A. scandens. As a matter of fact, it differs 
only in having very obtuse rather than acute leaves. The Quecchi 
name "cux" signifies "ear of corn." The ripe berries often are eaten, 
but only in small amounts and casually. 

Anthurium Scherzerianum Schott, Oesterr. Bot. Wochenbl. 
53. 1857; Regel, Gartenfl. pi. 482. 1865; Fl. Serres pi. 1794, 1795. 
1867-68; pi. 2454, 2455. 1880-83. Bot. Mag. pi. 5319. 1862. Type 
said to have been collected in Guatemala by Scherzer. No wild 
Guatemalan plants have been seen, and it may be that the type was 
collected in Costa Rica, where the species is common. 

Plants acaulescent or nearly so; petioles equaling or longer than the blades, 
10-25 cm. long; blades oblong-elliptic or oblong-lanceolate, 15-20 cm. long, 3-5 cm. 
wide, acuminate or cuspidate-acuminate, obtuse or rounded at the base, the pri- 
mary costal nerves numerous and subparallel, spreading, united to form a collective 
nerve remote from the margin; spathes subcoriaceous, bright red, broadly ovate, 
short-cuspidate, rounded and clasping at the base, 5-7 cm. long, 4-5 cm. wide; 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 317 

spadix subsessile, red, spirally twisted, 5-9 cm. long, 4 mm. thick; berries sub- 
globose, bright red. 

Called "lengua del diablo" in Costa Rica. This is, in Costa Rica, 
usually a terrestrial plant, growing in wet forest. It has long been a 
favorite plant in cultivation in Europe and the United States, where 
it has been hybridized with other species. Engler lists about 40 forms 
that are recognized in cultivation, distinguishable by the form of the 
leaves, color of the inflorescence, etc. In the wild state the inflor- 
escence is always bright red, but some of the cultivated forms have 
white or yellow spathes. The plant is so showy because of the bril- 
liantly colored inflorescences that it is hard to explain why it has not 
been collected recently in Guatemala, if it really occurs there, which 
we consider improbable. 

Anthurium Schlechtendalii Kunth, Enum. PI. 3: 75. 1841. 

Epiphytic in moist forest, 1200-1500 meters; Huehuetenango 
(Paso del Boqueron, along Rio Trapichillo below La Libertad, Stey- 
ermark 51171); San Marcos (above Finca El Porvenir, Volcan Taju- 
mulco, Steyermark 37078}. Southern Mexico; El Salvador; Costa 
Rica; Panama. 

Plants epiphytic or sometimes growing on rocks, rather large and coarse, the 
caudex very short and thick; petioles subtetragonous, less than one-fourth as long 
as the blade, usually very short, sometimes as much as 15 cm. long and 1-1.5 cm. 
thick, with a short node somewhat longer than broad; leaf blades obovate-oblong 
or oblanceolate-oblong, commonly 50-60 cm. long and 15-20 cm. wide, sometimes 
smaller or larger, acute or acuminate, coriaceous when dried, gradually narrowed 
from about the middle to the narrow cuneate base, the costa stout and prominent, 
sometimes 1 cm. thick at the base, the primary lateral nerves about 14 on each side, 
stout and prominent, not forming a distinct collective nerve, ascending at an angle 
of about 45 degrees; peduncles shorter than the leaves, often very short, sometimes 
60 cm. long, stout or slender; spathe reflexed, 10-25 cm. long or more, 2 cm. wide 
near the base, pale green or sometimes tinged with red or purple, long-attenuate, 
more or less decurrent at the base; spadix 10-25 cm. long, often 1 cm. or more in 
diameter at the base, attenuate upward, reddish or greenish, in fruit 2 cm. thick. 

Anthurium scolopendrinum (Ham.) Kunth, Enum. PI. 3: 68. 
1841. Pothos scolopendrinus Ham. Prodr. 16. 1825. A. longipes 
Matuda, Soc. Bot. Mex. 14: 23. 1952, not A. longipes N. E. Br. Card. 
Chron. 18: 297. 1882. 

On trees in moist or wet lowland forest, 1200 meters or lower; 
Izabal; Escuintla. British Honduras (Stann Creek District, Gentle 
2796, type of A. longipes Matuda, not N. E. Brown); Honduras to 
Costa Rica and Panama; South America. 



318 FIELDIANA: BOTANY, VOLUME 24 

Plants small, epiphytic, acaulescent or practically so, the caudex very short 
and rather thick; petioles slender, half as long as the blades or longer; leaf blades 
lanceolate, lance-oblong, or usually oblanceolate, thin when dried, acuminate, 
cuneate-attenuate to the base, 11-30 cm. long and 2-5 cm. wide, the costa slender, 
the primary lateral nerves numerous, oblique, ascending at an angle of about 45 
degrees, united to form a slender collective nerve close to the margin; peduncles 
very slender, equaling or exceeding the leaves; spathe narrowly lanceolate, green, 
much shorter than the spadix, 2.5-4 cm. long or shorter, subulate-attenuate; 
spadix very slender, sessile or subsessile, green, sometimes 12 cm. long but usually 
much shorter, obtuse; berries scarlet, 7 mm. long or less, few and scattered in the 
fruiting spadix. 

Illustrated, Pflanzenreich IV. 23B: /. 25. The characters by 
which Matuda attempts to separate his A. longipes, not A. longipes 
N. E. Brown, from A. scolopendrinum do not appear constant, and it 
seems best to consider the material studied as intergradients within 
a variable species. 

Anthurium seamayense Standl. Field Mus. Bot. 22: 4. 1940. 

Epiphytic or sometimes terrestrial on limestone in wet forest, 
2000 meters or lower; endemic; Alta Verapaz (type from Seraxcaj, 
C. J. Wilson 211}; Izabal; Huehuetenango (near Barillas). 

Plants acaulescent or nearly so, the caudex usually about 1.5 cm. thick, cov- 
ered with the fibrous remains of the cataphylls; petioles slender, 20-30 cm. long, 
geniculate about 1 cm. below the apex; leaf blades oblong-ovate to triangular- 
hastate or sometimes oblong-cordate or lance-oblong, mostly 20-40 cm. long and 
10-20 cm. wide near the base, acuminate or abruptly cuspidate-acuminate, deeply 
cordate at the base, with a very narrow or rather open sinus, the basal lobes some- 
times slightly overlapping, rounded, slightly extended inward or sometimes di- 
rected outward, the blade 5-nerved from the base, the primary costal nerves about 
8 on each side, united to form a collective nerve close to the margin; peduncles 
about equaling the petioles, very slender; spathe reflexed, lance-oblong or linear- 
lanceolate, 4.5-6 cm. long, 1 cm. wide or narrower, rather long-acuminate, brown, 
often twisted; spadix sessile, chocolate-brown, slender, 4-8 cm. long, 4-5 mm. 
thick, scarcely attenuate upward. 

The rather ample and excellently prepared material of this species 
recently collected shows considerable variation, and ultimately it 
may be broken down into two or three species. In the type the leaf 
blades are ovate-cordate with somewhat incurved basal lobes, while 
in several collections they are triangular-hastate with the basal lobes 
directed outward. There are no obvious concomitant characters for 
separating these forms. The extreme variant is Steyermark J^.1956 
from Izabal, in which the blades are narrowly lanceolate and about 
5.5 cm. wide. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 319 

Anthurium Seleri Engler, Bot. Jahrb. 25: 459. 1898. Nalastoc. 

On rocks, 800-1200 meters; endemic; Huehuetenango (type col- 
lected on limestone ruins at Chacala, Seler 2643; also between Nen- 
ton and Las Palmas). 

Plants acaulescent, with a short thick rootstock bearing many coarse brown 
fibers; petioles stout, half as long as the blades or longer, 10-15 cm. long, or some- 
times not more than 6 cm. long; leaf blades coriaceous when dried, yellowish green, 
pale or glaucescent beneath, narrowly elongate-sagittate, 20-35 cm. long, 5-7 cm. 
wide at the base, gradually narrowed to the long-attenuate apex, truncate or sub- 
cordate at the base, the basal lobes very short and rounded, the primary lateral 
nerves 8-9 on each side, ascending at a narrow angle, united to form a collective 
nerve remote from the margin; peduncles half as long as the leaves or longer; spathe 
clasping at the base, reflexed, dull green, lanceolate, equaling the spadix, 4-7 cm. 
long, 1.5 cm. wide or narrower, narrowly acuminate; spadix thick, scarcely atten- 
uate upward, 6-7 cm. long; berries oblong-obovoid, 6-7 mm. long. 

A well-marked species, related to A. parvispathum, easily recog- 
nized by the small auricles and the long and very narrow leaves. 

Anthurium silvigaudens Standl. & Steyerm. Field Mus. Bot. 
22: 69. 1940. Xacpec (Coban, Quecchi). 

Moist or wet forest, 1200-2000 meters; endemic; Alta Verapaz 
(type collected near Tactic, Standley 70505} ; Huehuetenango (Sierra 
de los Cuchumatanes) . 

Plants epiphytic or often terrestrial, sometimes growing on rocks, about a 
meter high, the short caudex about 1 cm. in diameter; petioles rather stout, about 
27 cm. long; leaf blades firmly membranaceous, ovate-cordate, about 35 cm. long 
and 22 cm. wide, cuspidate-acuminate, deeply (about 11 cm.) cordate at the base, 
the basal lobes broadly rounded, directed inward, the sinus closed, the basal nerves 
7, the lowest one emitting about 8 nerves within the basal lobe, the primary costal 
nerves about 8 on each side, extending almost or quite to the margin; peduncles 
short, 7 cm. long; spathe membranaceous and thin when dried, pale green, ovate 
or lance-ovate, 8-11 cm. long and 3-5 cm. wide, long-acuminate, wavy-margined, 
broadly clasping and cordate at the base, conspicuously nerved; spadix subsessile, 
dull purplish green, 1 cm. thick. 

Although found growing on the ground in the wet Tactic forest, 
it has also been found as an epiphyte in the Sierra de los Cuchuma- 
tanes. It often is grown for ornament in the gardens of Coban. 

Anthurium subcordatum Schott, Oesterr. Bot. Zeitschr. 181. 
1858. A. Coibionii Standl. & Steyerm. Field Mus. Bot. 23: 209. 1947 
(type collected in Sierra de las Minas, Zacapa, Steyermark 42212). 

Moist or wet forest, 1100-2500 meters; endemic; Alta Verapaz; 
Zacapa; Chiquimula; El Progreso; Jalapa; Quiche"; Quezaltenango 



320 FIELDIANA: BOTANY, VOLUME 24 

(type of A. subcordatum collected from Las Nubes on the slopes of 
Volcan de Zunil, Wendland) ; Huehuetenango. El Salvador. 

Plants epiphytic or terrestrial, acaulescent or nearly so, the caudex short and 
thick; petioles slender or stout, 13-35 cm. long; leaf blades chartaceous or perga- 
mentaceous to coriaceous when dried, coriaceous or fleshy when alive, oblong or 
ovate-oblong, 15-60 cm. long, as much as 25 cm. wide, acute or acuminate, broadly 
rounded or subcordate at the base and 5-7-nerved, the primary costal nerves 6-10 
on each side, united to form a collective nerve remote from the margin; peduncles 
12-50 cm. long, slender; spathe ascending to reflexed, oblong-lanceolate or linear- 
lanceolate, 4-6 cm. long, 2 cm. wide, long-cuspidate, short-decurrent at the base, 
green; spadix borne on a stipe as much as 3 cm. long in fruit, 5-10 cm. long and 
0.5-2 cm. thick, green to dull lavender, becoming red in fruit. 

A Guatemalan collection of this species (Heyde & Lux 3516) was 
once listed by Captain Smith as A. quinquenervium Kunth, a closely 
related Colombian species, with which it may eventually have to be 
united. Engler separates the two on a slight difference in leaf shape 
and the relative width at the base, and it is suspected that such differ- 
ences will prove to be unreliable in maintaining the two species apart. 

Anthurium tetragonum Hook, ex Schott, Prodr. Aroid. 475. 
1860. 

Epiphytic or terrestrial, often growing on rocks, 1100 meters or 
lower; Alta Verapaz; Izabal; Zacapa. Southern Mexico; British 
Honduras to Panama. 

Plants large and coarse, acaulescent or nearly so, the short caudex often 6-8 cm. 
thick; petioles scarcely one-sixth as long as the blades, commonly 10-20 cm. long 
or shorter, broadly canaliculate above, flat beneath, the short node thick and about 
as broad as long or broader; leaf blades coriaceous when dried, broadly obovate to 
oblanceolate, 60-130 cm. long and 25-50 cm. wide when well developed, rounded 
and short-cuspidate at the apex, cuneately narrowed to the base, the costa very 
thick, subpentagonal, usually 1-2 cm. thick at the base, the primary lateral nerves 
15-20 on each side, erect-spreading, not united to form a distinct collective nerve, 
stout and prominent; peduncles terete, usually one-third as long as the blades or 
shorter, mostly 20-30 cm. long; spathe oblong-lanceolate, long-cuspidate, obliquely 
decurrent at the base, 15-20 cm. long, 3-4 cm. wide near the base; spadix short- 
stipitate, 2-3 cm. thick at the base, caudate-attenuate upward, dull green or brown- 
ish or dark purplish, 15-25 cm. long; sepals 3 mm. long and one-third as wide; 
stamens equaling the sepals; ovary subfusiform, half longer than the sepals, atten- 
uate to the apex; berries elongate-turbinate, purplish below, yellowish green toward 
the apex, 1-1.5 cm. long, 2-3 mm. thick, 1-3-seeded; seed oblong, one-third as long 
as the berry. 

Called "hoja de viento" in Veracruz. This species and A. crassi- 
nervium are much alike in general appearance, and in herbarium 
specimens are sometimes difficult to distinguish, but the live plants 
can be separated easily by the very different petioles, which in A. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 321 

tetragonum have a flat node not carinate dorsally, whereas in A. cras- 
sinervium the node is dorsally 3-carinate. 

Anthurium titanium Standl. & Steyerm. Field Mus. Bot. 23: 
211. 1947. 

Known only from the type, terrestrial in moist or wet forest, San 
Marcos, Volcan de Tajumulco, above Finca El Porvenir, along Rio 
Cabus, 1300-1500 meters, Steyermark 37963. 

Plants very large and coarse; petioles 55 cm. long or probably much longer in 
adult leaves, scarcely vaginate below but slightly dilated, geniculate about 1.5 cm. 
below the base of the blade, the node 1.5 cm. broad; leaf blades chartaceous or 
thick-membranaceous when dried, rounded-cordate, 50-120 cm. long, about 40-60 
cm. wide or probably even wider, obtuse or rounded at the apex and broadly short- 
cuspidate, deeply cordate at the base with a very broad and open sinus, entire, 
7-nerved from the base, the lowest 2 nerves almost 1 cm. broad at the base, naked 
on the outer margin for 4-6 cm., emitting 5-6 stout nerves on the lower side, the 
primary lateral nerves slender, remote, prominent, forming an irregular collective 
nerve remote from the margin, the veins prominulous, laxly reticulate; peduncle 
70 cm. long or more, stout, 2 cm. thick near the base; spathe coriaceous, erect, 
green suffused with red, lanceolate, 20-26 cm. long, 4.5-6.5 cm. wide near the base, 
very narrowly attenuate-acuminate, strongly clasping at the base; spadix bronze or 
brownish green, caudiform, about 19-30 cm. long, 12-18 mm. thick near the base 
in the dried state, in the living state as much as 2.5-5 cm. thick below, gradually 
attenuate upward, longer than the spathe, the stout stipe 3-4.5 cm. long, as much 
as 12 mm. thick. 

Anthurium Tuerckheimii Engler, Bot. Jahrb. 25: 380. 1898; 
Pflanzenreich IV. 23B:/. 30, F-L. 1905. 

Epiphytic in wet forest, 350-1650 meters; Alta Verapaz (type 
from Pansamala, Tuerckheim 864); Quezaltenango. British Hon- 
duras; Honduras. 

Caudex slender, generally much elongate, with often greatly elongate inter- 
nodes, sometimes 1 cm. thick but usually much more slender; petioles slender, 
equaling or more often shorter than the blades; leaf blades thin and almost mem- 
branaceous when dried, oblong-elliptic or broadly lance-oblong, 18-25 cm. long, 
7-10 cm. wide, acuminate or cuspidate-acuminate, broadly rounded at the base 
and often emarginate or subcordate, the primary costal nerves 8-10 on each side, 
slender, united to form a slender connective nerve remote from the margin; pedun- 
cles very slender, 12-15 cm. long; spathe linear-lanceolate, green, about 5 cm. long 
and 8 mm. wide, narrowly long-acuminate; spadix subsessile, yellowish green, ob- 
tuse, 3.5-6 cm. long, 5-8 mm. thick. 

Anthurium verapazense Engler, Pflanzenreich IV. 23B: 191. 
1905. 



322 FIELDIANA: BOTANY, VOLUME 24 

Wet forest, epiphytic or terrestrial, sometimes on limestone cliffs, 
350 meters or lower; Alta Verapaz (type from Cubilguitz, Tuerck- 
heim 7788}; Izabal. British Honduras. 

Plants acaulescent or nearly so, the caudex short, about 1.5 cm. thick; petioles 
about 45 cm. long, vaginate at the base for 6-7 cm., slender; leaf blades subcori- 
aceous when dried, elongate-sagittate or deltoid-sagittate, about 40-50 cm. long 
and 15-22 cm. wide but sometimes larger or smaller, the terminal lobes elongate- 
triangular, apiculate or cuspidate, the basal lobes broadly oblong, rounded at the 
apex, slightly directed outward, separated by an oblong or sometimes triangular 
sinus about 7-13 cm. deep, the primary costal nerves 10 or more on each side, 
slender, united to form a collective nerve close to the margin; peduncles 70 cm. 
long or shorter, slender; spathe linear-lanceolate, 6-15 cm. long and 4-12 mm. 
wide or shorter, long-acuminate, green or brownish green; spadix sessile, very slen- 
der, in anthesis 12-25 cm. long and 4-8 mm. wide, in fruit sometimes longer and 
thicker; berries obovoid, 5 mm. long, 4 mm. thick, orange or yellow. 

This has been reported from British Honduras as A. concinnatum 
Schott. 

CALADIUM Ventenat 
Reference: Engler, Pflanzenreich IV. 23E: 23. 1920. 

Terrestrial plants, arising from tuberous rhizomes; leaves long-petiolate, the 
blade usually peltate and sagittate, often colored; peduncles usually long and soli- 
tary; tube of the spathe convolute, persistent, constricted at the apex, the blade 
cymbiform, whitish; spadix slightly shorter than the spathe, naked and stipitate at 
the base, the pistillate lower portion cylindric-conoid or ellipsoid, densely many- 
flowered, the sterile staminate portion of the spadix subconic, longer than the pistil- 
late, the fertile staminate portion contiguous with the sterile, subclavate, densely 
flowered, twice as long as the pistillate part; flowers unisexual, naked; staminate 
flowers with 3-5 stamens, these connate to form a truncate-obpyramidal synan- 
drium sinuately subhexagonal at the apex, the connective thick, the anther cells 
opening by a short, apical slit; ovary of the pistillate flower usually 2-celled, with 
several ovules in each cell; stigma depressed-hemispheric, slightly 3-4-sulcate; fruit 
baccate, several-many-seeded; seeds ovoid. 

A genus of about 16 species, all natives of South America, chiefly 
of Brazil. 

Caladium bicolor (Ait.) Vent. Descr. Gels. pi. 30. 1800. Arum 
bicolor Ait. Hort. Kew. 3 : 316. 1789. Hoja de leche; Corazon sangri- 
enta; Corazon de Maripa; Quequeshque manchado; Capa de rey; Hoja 
de adorno. 

Planted commonly in Guatemalan gardens and sometimes more 
or less naturalized about dwellings or in cemeteries in the moister 
lowland regions. Native probably of the Amazon Valley. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 323 

The plant is easily recognized by its peltate leaves, which always are spotted 
or striped with white, cream, pink, and other colors. The plants show infinite 
variety in leaf coloring. 

This species is much cultivated for ornament in Europe and the 
United States because of its striking and beautiful leaves. It is seen 
frequently also in gardens of the uplands in Guatemala, but does not 
thrive there so well as in the tierra caliente. 



COLOCASIA Schott 

Reference: Krause, Pflanzenreich IV. 23E: 62. 1920. 

Plants terrestrial, usually acaulescent and with tuberous roots; petioles long- 
vaginate; blades often very large, peltate, ovate-cordate or sagittate-cordate, the 
posterior lobes rounded; peduncles usually several; tube of the spathe one-fifth to 
one-half as long as the blade, ovoid or oblong, convolute, persistent and somewhat 
accrescent in fruit, the blade oblong to lanceolate, finally separating from the tube; 
spadix shorter than the spathe, the pistillate portion short, the sterile staminate 
portion short and narrow, the fertile staminate part long and slender, the short 
apex of the spadix sterile; flowers unisexual, naked; staminate flowers with 3-6 
stamens, these connate to form an obpyramidal synandrium dilated, subtruncate, 
and irregularly polygonous at the apex; anther cells linear or linear-oblong, opening 
by a very short, apical slit; pistillate flowers 3-4-gynous, the ovary ovoid or oblong, 
1-celled, the ovules several or numerous; stigma sessile or nearly so, depressed- 
capitate, shallowly 3-5-sulcate; berries obconic or oblong, greenish, many-seeded; 
seeds oblong, with abundant endosperm. 

Seven species are known, natives of tropical Asia. 

Colocasia esculenta (L.) Schott, Melet. 1 : 18. 1832. Arum escu- 
lentum L. Sp. PI. 965. 1753. A. Colocasia L. loc. cit. C. antiquorum 
Schott, Melet. 1: 18. 1832. Coco (name used by negroes of North 
Coast, probably African in origin); Xacox (Coban, Quecchi); Ma- 
langa; Papa malanga; Elephant-ear; Dasheen. 

Planted in various regions of Guatemala for ornament and occa- 
sionally in the lowlands for food; more or less naturalized in the 
North Coast and Alta Verapaz. Native of the East Indies but now 
grown generally in tropical and subtropical regions. 

Plants very variable in size, the thick petiole often a meter long, the blades 
50 cm. long or in the ornamental forms even much larger, peltate-ovate, cordate at 
the base, the anterior lobe broadly ovate, abruptly short-pointed; peduncles much 
shorter than the petioles; spathes pale yellowish, 15-35 cm. long, the limb lanceo- 
late, 2-5 times as long as the tube. 

The ornamental form of this species is not especially common in 
Central American gardens, but it is seen now and then. In the United 
States this form is known by the name "elephant-ear," or more often 



324 FIELDIANA: BOTANY, VOLUME 24 

as "caladium," the latter given because the plant is confused with 
the genus Caladium. The form planted for its rounded, edible tubers 
is called "dasheen" in English-speaking regions, in the West Indies 
"malanga" and "yautia." The dasheen is planted for food in lower 
regions of the north coast and Alta Verapaz, as in the region of Car- 
cha. The tubers are poisonous or at least very dangerous if eaten 
raw, and must be prepared for the table by long exposure to heat, 
preferably by boiling first with soda or some other agent that will 
neutralize and destroy the calcium oxalate crystals. When prop- 
erly cooked, the tubers are of good flavor and agreeable to eat, with 
a rather waxy consistency and nutty taste. Attempts have been 
made to introduce the dasheen into the Gulf Coast area of the United 
States, but with little success. It does grow and produce abundantly, 
but the people apparently do not like it as a food. The Maya name 
is reported from Yucatan as "cucutmacal." 



DIEFFENBAGHIA Schott 

Reference: Engler, Pflanzenreich IV. 23Dc: 36. 1915. 

Stout, terrestrial herbs with white or yellow sap, the stems often much elongate 
and prostrate and rooting, or the plants almost acaulescent, leafy at the apex of the 
caudex; leaves thick and succulent, long-petiolate, the petiole vaginate to the mid- 
dle or higher, terete near the apex, the blades oblong, with a thick costa, the pri- 
mary nerves numerous, parallel with the secondary ones, arcuate near the apex; 
peduncles shorter than the leaves, the spathe oblong, persistent, the lower portion 
convolute, the throat open and expanded into a straight or recurved blade; spadix 
erect, slightly shorter than the spathe, the lower portion densely many-flowered, 
pistillate, dorsally adnate to the spathe, the upper staminate portion free, subcy- 
lindric, densely many-flowered, separated from the pistillate portion by an almost 
naked portion bearing a few abortive staminate flowers; flowers by abortion uni- 
sexual, naked, the staminate with 4-5 stamens, these connate to form a sessile, 
thickened, truncate, apically 4-5-sulcate synandrium; pistillate flowers with 4-5 
claviform staminodia, the pistil 2-3-carpellate, sessile, depressed-ovoid, the thick 
carpels concrete along one side; ovules solitary in the cells, erect; style none, the 
stigma 2-3-lobate, concave in the middle, with thick lobes; fruit baccate, crowned 
by the remains of the stigmas, 1-3-celled; seeds globose or ovoid. 

About 25 species are known, generally distributed in tropical 
America. They grow chiefly in wet, lowland forests. Some species 
long ago were introduced into cultivation in greenhouses of Europe 
and the United States, and they are often seen, especially those forms 
with leaves variegated in white or yellow, as house plants in the 
United States. The plants with variegated leaves are apparently 
only variants of the normal green-leaved plants, and often are found 
in the forest. Six species of Dieffenbachia are recorded from Central 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 325 

America, chiefly in Costa Rica and Panama. The plants are well 
known to people living in the areas where they grow. When cut, 
they exhale an unpleasant mephitic odor. The sap is said to be very 
irritating to the skin, and to cause serious inflammation, but its repu- 
tation is exaggerated. An English name for the plants is "dumb 
cane," and in the United States the name "mother-in-law plant" is 
sometimes applied to them (this name, rather strangely and prob- 
ably because of some confusion with Dieffenbachia, is now applied 
more commonly in the United States to Sansevieria) . In Panama 
they are called "oto de lagarto," "ot6" being the term applied to the 
cultivated Xanthosomas. 

Leaves conspicuously spotted with white or cream D. picta 

Leaves green. 

Spathe orange at maturity D. Oerstedii 

Spathe green or yellowish green at maturity. 

Sheath of the petiole extending almost to the base of the leaf blade . .D. Pittieri 
Sheath of the petiole ending far below the blade D. seguina 

Dieffenbachia Oerstedii Schott, Oesterr. Bot. Zeitschr. 8: 179. 
1858. Flor de zahino; Colch (San Marcos); Cuyanigua. 

Frequent in wet, lowland forests of both coasts, ascending com- 
monly to 750 meters or rarely to as much as 1900 meters; Izabal; 
Jutiapa; Santa Rosa; Escuintla; Suchitepe"quez; Retalhuleu; San 
Marcos. Ranging southward to Panama. 

Plants usually about a meter high, and with erect stems; petioles 12-20 cm. 
long, vaginate for two-thirds or more of their length, slender above the sheath, the 
blades oblong-ovate or broadly oblong, 12-30 cm. long and 6-12 cm. wide, or even 
larger, very thick when fresh but drying thin, short-cuspidate or acute, rounded or 
subcordate at the base; peduncles about 7 cm. long, the spathe 15 cm. long and 
3-4 cm. wide, subacute, becoming bright orange when mature; spadix slightly 
shorter than the spathe, naked at the base, the pistillate portion 5 cm. long, the 
stamina te of the same length, the two separated by a sterile portion 2 cm. long; 
ripe berries scarlet. 

Called "hoja de puerco" on the Atlantic coast of Honduras. In 
favorable localities, especially in the Pacific lowlands, the plants, 
about a meter high, form extensive and dense colonies. 

According to Birdsey, present authority on the Araceae, this and 
the following species should probably be merged as a single poly- 
morphic species. Since Dieffenbachia picta is the earlier available 
name, it would become the proper name if the two species are 
united. 



326 FIELDIANA: BOTANY, VOLUME 24 

Dieffenbachia picta (Lodd.) Schott, Oesterr. Bot. Wochenbl. 
2: 68. 1852. Caladium pictum Lodd. Bot. Cab. pi. 608. 1822. Que- 
quesque; Acacia (Jutiapa, the latter a rather fantastic name). 

Common in cultivation for ornament in Guatemalan gardens; 
sometimes found also in the lowland forests. 

Similar to the ordinary native species, and distinguishable only by the varie- 
gated leaves, which are green and variegated or striped with white or cream. 

By Engler this is called a "typus polymorphus," and very justly, 
since there can be no doubt that the "species" consists of variegated 
forms of various species. These forms have long been cultivated in 
hothouses of Europe, and are also favorite pot plants in the United 
States, where they thrive well under conditions of heat and dryness. 
It is probable that the plants noted in Guatemalan gardens have been 
brought from the forests, and are mere forms of the local species. 

Dieffenbachia Pittieri Engl. & Krause, Pflanzenreich IV. 23Dc: 
42. 1915. 

Wet forest at sea level; Izabal. British Honduras; Atlantic coast 
of Honduras; Panama. 

Plants about a meter high or less, with thick stems as much as 60 cm. tall; 
petioles mostly 10-13 cm. long, white at the base; leaf blades obliquely oblong, 
18-35 cm. long, 6.5-19 cm. wide, acuminate, obtuse or subacute at the base, dark 
green above, paler beneath; peduncles 4-7 cm. long, the spathe 16-20 cm. long or 
more, long-acuminate, pale green; staminate portion of the spadix 6-7 cm. long, 
separated from the fertile portion by a sterile part 3 cm. long. 

Called "dumb cane" in British Honduras, "puerco" and "quisca- 
mote" in Honduras. 

Dieffenbachia seguina (L.) Schott, Melet. 1: 20. 1832. Arum 
seguinum L. Sp. PI. ed. 2. 1371. 1763. Coche de monte (fide Morales) ; 
Cuyamiz; Hierba de sapo (fide Aguilar). Figure 55. 

Damp or wet forest, chiefly in the lowlands, rarely ascending to 
1200 meters; Pete"n; Alta Verapaz; Santa Rosa; Escuintla; Guate- 
mala; Sacatepquez; Chimaltenango; Solola; Suchitepe"quez; Retal- 
huleu; Huehuetenango; San Marcos. Southern Mexico (Chiapas); 
El Salvador; Honduras to Costa Rica; West Indies; South America. 



FIG. 55. Dieffenbachia seguina. a, Flower with spathe and spadix (X 
b, Spadix (X Yz). c, Pistillate flower (X 8). d, Leafy apex of shoot (X 
e, Staminate flower (X 7). 




327 



328 FIELDIANA: BOTANY, VOLUME 24 

Plants about a meter high, the caudex usually thick and often decumbent, 
4-5 cm. thick; petioles 40 cm. long or less, the sheath half as long or longer, the 
blades oblong to oblong-ovate, as much as 75 cm. long and 22 cm. wide, dark green 
above, slightly paler beneath; spathes as much as 27 cm. long, the closed portion 
15 cm. long and 3 cm. broad, the upper portion 10 cm. long, with a cusp 2 cm. long; 
basal portion of the spadix naked, 2 cm. long, the densely flowered pistillate portion 
9 cm. long, the staminate portion 9 cm. long, separated from the pistillate by a 
naked portion 2 cm. long; berries scarlet. 

Engler refers the North American material to var. viridis Engler 
of D. seguina, which he calls a "polymorphic type/' but the various 
varieties and subvarieties that he recognizes seem to have little syste- 
matic importance, and are of concern chiefly to horticulturists. Called 
"hoja de puerco" in Honduras and "cuyanigua" in El Salvador. 



MONSTER A Adanson 

Reference: Engler & Krause, Pflanzenreich IV. 23B: 97. 1908. 

Scandent epiphytes, usually with thick and greatly elongate stems, these root- 
ing at the nodes; leaves distichous, those of juvenile plants small, ovate or broadly 
ovate-cordate, short-petiolate, appressed to tree trunks and covering the stem and 
roots; petiole vaginate to the middle or higher, the blades variable, usually some- 
what unequilateral, broadly oblong or oblong-ovate, entire or frequently perforated 
with large holes, sometimes pinnatifid; peduncles solitary or several at the tips of 
the branches, the spathe ovate or oblong-ovate, apiculate, cymbiform-convolute, 
closed after fecundation of the flowers and finally deciduous; spadix sessile, free, 
subcylindric, densely many-flowered, somewhat shorter than the spathe, the lowest 
flowers sterile, the others perfect; fertile flowers with 4 stamens, the filaments rather 
broad, somewhat compressed, abruptly narrowed at the apex into the slender, 
acuminate connective; anthers 2-celled, the cells oblong, exceeding the connective; 
ovary obconic-prismatic, 2-celled, the ovules 2 in each cell; style equaling the ovary, 
truncate at the apex, the stigma depressed-oblong or linear; fruits baccate, crowded, 
the disklike terminal portion soon freed from the rest of the berry, 1-3-seeded ; 
seeds obovoid or subcordate, subcompressed. 

About 27 species are known, widely dispersed in tropical America. 
The plants are abundant in many of the Guatemalan forests, espe- 
cially in the more humid parts of the tierra caliente. Because of their 
great size and their striking foliage, they constitute a conspicuous 
part of the epiphytic vegetation of the taller trees. Most of the 
plants have distinctive leaves, perforated with large holes, a feature 
found in only one other Central American group of plants (palms of 
the genus Reinhardtia) . 

The aerial roots of these plants, as well as those of some other 
aroids, particularly Philodendron, often attain a great length, reach- 
ing from the branches of fairly high trees almost or quite to the 
ground. They are much used in Guatemala for making the so-called 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 329 

mimbre furniture, similar to the light rattan furniture made com- 
monly in the United States. The dried roots, of uniform diameter, 
or sometimes the fresh ones, are wound tightly and evenly about a 
wooden frame, forming handsome and durable articles of furniture. 
The fruiting spadices of some of the species, particularly M. deli- 
ciosa, become whitish at maturity and very juicy. They are sweet 
and of good flavor and often are eaten, but care must be taken to 
eat only fruits that are thoroughly ripe, since immature ones will 
cause swelling and irritation of the mouth because of needle-like 
crystals of calcium oxalate found in the tissues or sap. Because of 
the unique leaves, these plants long ago were introduced into culti- 
vation in hothouses of Europe and the United States. They have 
become common house plants in some localities because their thick 
and fleshy leaves withstand well the drying effects of ordinary steam- 
heated air. 

Leaves, even the adult ones, entire and without perforations, or rarely with a very 

few, inconspicuous, irregularly scattered perforations. 

Leaves (some) with a few small inconspicuous perforations, some without per- 
forations. Leaf blades rounded at the base, 30-40 cm. long, 13-20 cm. wide. 

M . belizensis 
Leaves all without perforations. 

Leaf blades obtuse or rounded at the base M. acuminata 

Leaf blades cordate or subcordate at the base or sometimes acute. 

Mature leaves mostly 10-15 cm. long, on petioles 3.5 cm. long or shorter; 
spadices strongly tuberculate because of the much elongate, conic 

styles M. tuberculata 

Mature leaves about 50 cm. long and 25 cm. wide, on petioles about 25-35 
cm. long; spadices not tuberculate, the styles very short. 

M. grandifolia 

Mature leaves perforated or pinnatifid, or both, the perforations usually numer- 
ous, sometimes few but then large and conspicuous. 

Mature leaves regularly pinnatifid, conspicuously cordate M. deliciosa 

Mature leaves perforated, the margins sometimes irregularly cleft but never 

regularly pinnatifid, acute to cordate at base. 

Perforations of the mature leaves irregularly scattered or rarely arranged in 
a single series, sometimes irregularly cleft; leaves acute to rounded at 

base; from sea level to 1100 meters M. perttisa 

Perforations of the mature leaves arranged in 2-4 more or less regular rows 
along the costa of the blade; leaves rounded to cordate at base; gener- 
ally occurring between 1300 and 2700 meters M. Friedrichsthalii 

Monstera acuminata C. Koch, Ind. Sem. Hort. Berol. App. 4. 
1855 (described from plants cultivated in Berlin, obtained somewhere 
in Central America by Warscewicz). Sehuanen (Alta Verapaz). 

Epiphytic on trees in moist or wet forest, 1000 meters or lower, 
most common at or little above sea level; Alta Verapaz; Izabal; 



330 FIELDIANA: BOTANY, VOLUME 24 

Santa Rosa; Escuintla; Huehuetenango. Costa Rica; reported from 
Amazonian Brazil. 

A small or large vine with thick stems; petioles of adult leaves 7.5-24 cm. long, 
the blades rather thick, oblong-ovate, mostly 20-38 cm. long and 10-23 cm. wide 
but somewhat variable in size, very asymmetric, acute or obtuse and usually short- 
cuspidate, obliquely rounded at the apex; peduncles short, often very short; closed 
spathe white, oblong, 7-18 cm. long, with an acumen 1 cm. long; spadix oblong- 
ovoid or oblong, 5-16 cm. long and 1.5-3 cm. thick, or larger; pistils before maturity 
ovoid, gradually narrowed into the short stigma. 

Monstera belizensis Lundell, Lloydia 2: 76. 1939. 

Known only from the type, British Honduras, wet lowland forest, 
Valentin, El Cayo District, C. L. Lundell 6198. 

A large epiphytic vine; leaves large, the petioles 22-27 cm. long, vaginate to 
the middle of the upper node, this 1.5-2.5 cm. long; leaf blades thin, obliquely 
ovate-oblong or lance-oblong, 32-44 cm. long, 13-20 cm. wide, narrowed at the 
apex or rounded or obtuse and apiculate, rounded and unequal at the base, usually 
entire, sometimes with 1-2 small perforations remote from the costa, the principal 
lateral nerves 9-12 on each side; peduncles about 7 cm. long, 8-10 mm. thick; 
spathe unknown; fruiting spadix sessile, cylindric, 15 cm. long, 3.5 cm. thick; fruits 
cylindric, 7-9 mm. long (excluding the style), 1-2-seeded; seeds oblong-ellipsoid, 
slightly compressed laterally, 6.5-9 mm. long; style cylindric, 4-6 mm. long, trun- 
cate or slightly concave, the stigma plane. 

We have seen no material of this species. Until much more mate- 
rial is available for study, based upon herbarium as well as field re- 
search, it is not possible at the present time to be certain of the status 
of M. belizensis, M. acuminata, and M. grandifolia. It is possible 
that they represent variations of the same taxon. 

Monstera deliciosa Liebm. Dansk. Vid. Medd. Forh. 19. 1849- 
50. Pinanona; Harp6n. 

Moist or wet, mountain forest, 900-1500 meters; San Marcos; 
Huehuetenango; often grown for ornament in gardens, or as a house 
plant. Southern Mexico; Costa Rica; Panama. 

Usually a large and coarse, epiphytic vine, the stems stout, often greatly elon- 
gate, sometimes 6 cm. thick; petioles often a meter long and 2-2.5 cm. thick; blades 
of the earliest leaves small, cordate, entire, those of the succeeding leaves ovate- 
cordate, entire, or becoming pinnatifid; adult blades 40-60 cm. long or more, rather 
thick and somewhat coriaceous, lustrous and bright green above, paler beneath, 
regularly pinnatifid, the oldest ones also with numerous small perforations close to 
the costa, in the dried state the ultimate veins conspicuously lineolate-reticulate; 
peduncle 10-15 cm. long, the spathe coriaceous, broadly ovate and cymbiform, 
minutely apiculate, 20-25 cm. long, pale yellow; spadix cylindric, 11-20 cm. long, 
pale yellow or in fruit whitish; stigmas linear; berries 1 cm. long. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 331 

Sometimes called "ceriman" in the United States, the origin of 
that term unknown to the writers. This is the species most often 
seen in cultivation outside tropical America, and it probably pro- 
duces better edible fruits than other members of the genus. In 
Guatemala it has been found growing wild in only four localities. 
The deeply cordate leaves are quite characteristic of this species. 

The plant is often grown for its succulent fruits in tropical or sub- 
tropical areas where it is not native. Large plants of this species 
often are seen as pot or tub plants in the patios and corredores of 
Guatemala. Planted in the ground they often become a pest because 
of their rapid growth and their tendency to monopolize small spaces 
allotted to a variety of cultivated plants. The aerial roots are used 
in Mexico for making strong baskets. 

Plants from Costa Rica and Panama belonging to this species 
have sometimes been misidentified as M. dilacerata Koch. So far as 
known, M. dilacerata is a South American species that reaches its 
northern limit in Panama. 

Monstera Friedrichsthalii Schott, Oesterr. Bot. Wochenbl. 4: 
65. 1860 (based on material said to have been collected in Guatemala 
at "Niquinomo" by Friedrichsthal; this is probably not a Guate- 
malan locality but is perhaps Nicaragua). M. siltepecana Matuda, 
Rev. Soc. Mex. Hist. Nat. 11: 97. pi. 2, fig. 9. 1950 (type collected at 
Cascada, Siltepec, Chiapas, Matuda 18642). Hoja de colador. 

On trees in wet forests, generally between 1300 and 2700 meters; 
Alta Verapaz; Baja Verapaz; El Progreso; Zacapa; Jalapa; Santa 
Rosa; Escuintla; Guatemala; Sacatepe"quez; Chimaltenango; Re- 
talhuleu; Quezaltenango; San Marcos. Chiapas; El Salvador; Hon- 
duras; Costa Rica; Panama. 

A small or large vine; petioles 12-47 cm. long; leaf blades obliquely ovate, very 
asymmetric, one side often almost twice as wide as the other, acute or rather long- 
acuminate, rounded or cordate at the base, rather thin, the perforations small or 
large, 2-3-seriate with other small ones between the large ones; peduncles slender, 
terete, 7.5-10 cm. long, much shorter than the petiole; spathe 11-22 cm. long, 8-17 
cm. broad, greenish to whitish without, pink or rose within, deciduous; spadix 
cylindric, obtuse, 9-13 cm. long, 1.5-2.2 cm. thick; pistils subconic, about 1 cm. 
long, the sessile stigma linear-oblong; berries truncate. 

Called "pinanona" in El Salvador. For an illustration see Pflan- 
zenreich IV. 23B:/. 42. 1905. 

It is possible that this species and M. pertusa are variations of 
one taxon. In general, M. Friedrichsthalii occurs in Guatemala at 
much higher elevations than M. pertusa; M. pertusa usually occurs 



332 FIELD IANA: BOTANY, VOLUME 24 

from sea level to less than 1000 meters, whereas M. Friedrichsthalii 
occurs generally between 1300 and 2700 meters. Monstera siltepe- 
cana Matuda, described from Chiapas, cannot be separated from the 
many variations of leaf size and perforation manifested by numerous 
specimens of M. Friedrichsthalii. 

Monstera grandifolia Standl. & Steyerm. Field Mus. Bot. 23: 
212. 1947. 

Known certainly only from the type, Pete*n, low forest between 
Finca Yalpemech and Chinaja, 50-100 meters, Steyermark 45423; 
Izabal(?). 

A large coarse epiphytic vine, the stems almost 2 cm. thick; petioles slender, 
about 25-38 cm. long, narrowly vaginate throughout, the node about 15 mm. long 
and 8 mm. thick; leaf blades rather thick, oblong-ovate, 47-50 cm. long and 25-33 
cm. wide, or sometimes larger and as much as 35 cm. wide, apparently obtuse or 
rounded at the apex and short-cuspidate, somewhat narrowed below, the base sub- 
cordate and somewhat unequal, the costa thick and prominent beneath, the pri- 
mary lateral nerves coarse and prominent; peduncles about 11 cm. long and 1 cm. 
thick; spathe unknown; spadix cylindric, sessile, 18 cm. long, 3 cm. thick, the 
flowers very dense; style very short and broad, truncate or subtruncate. 

A collection from Izabal that probably represents the same spe- 
cies has leaf blades that are distinctly acute at the base. In other 
respects, especially in the size and texture of the leaves and the 
appearance of the spadix, it is like the type of M. grandifolia. 

Monstera pertusa (L.) de Vriese, Hort. Spaarn-Bergens. 40. 
1839. Dracontium pertusum L. Sp. PI. 968. 1753. M. longipeduncu- 
lata Matuda, Bol. Soc. Bot. Mex. 14: 22-23. fig. 1. 1952. Harpdn; 
Madre de harpdn; Colcuc (Quecchi). Figure 56. 

Moist or wet forest, chiefly at the lower altitudes from near sea 
level to 1100 meters, but sometimes found as high as 1500 meters; 
Pete"n; Alta Verapaz; Chiquimula; Santa Rosa; Escuintla; Suchite- 
pquez; Retalhuleu; Huehuetenango. Honduras to Panama; South 
America. 

A large or small, epiphytic vine, the stems 1-3 cm. thick; blades of juvenile 
leaves ovate or oblong-ovate or obliquely oblong-elliptic, without perforations; 



FIG. 56. Monstera pertusa. a, Portion of surface of fertile part of spadix 
(X 6). 6, Portion of surface of sterile part of spadix (X 6). c, Staminate flower 
(X 7). d, Pistillate flower from middle part of spadix (X 7). e, Portion of 
flowering shoot with leaves ( X M). /, Longitudinal section through pistil ( X 
g, Spadix with spathe (X ^). 




333 



334 FIELDIANA: BOTANY, VOLUME 24 

petioles of adult leaves up to 20-38 cm. long, the blades rather thick, broadly 
ovate-oblong, 30-60 cm. long and 15-26 cm. wide or sometimes larger, somewhat 
asymmetric, acute or rounded at the base, acute or short-acuminate at the apex, 
the perforations irregularly scattered or in a single series near the costa, usually 
large, often unequally and irregularly pinnatifid, the segments 1-5, broadly rhom- 
bic-oblong to linear-oblong, truncate, 10-17 cm. long, 2-7 cm. wide; peduncle 5-20 
cm. long, the spathe coriaceous, ovate, oblong, or broadly orbicular, acuminate, 
9-25 cm. long, 10-14 cm. wide, at first green, becoming whitish or yellowish; spadix 
cylindric, 6-10 cm. long and 1.5-2.5 cm. thick; pistils subprismatic, whitish to pale 
yellow, about 4 mm. long, the stigma suborbicular or broadly oblong; berries obo- 
void, with the style 5-6 mm. long, 4 mm. thick, mostly 1-seeded. 

Engler and Krause refer the Central American material of this 
species to var. Jacquinii (Schott) Engler, but this seems to differ in 
no essential respect from other forms of M. pertusa, which these auth- 
ors describe as a "polymorphic type." There is great variation in the 
size and number of perforations in the adult foliage, and in the tend- 
ency for these perforations to become enlarged; often a leaf splits 
apart and separates to form two or more irregular segments, giving 
the leaf an unequally pinnatifid appearance. Upon the latter type 
of leaf Matuda based his M. longipedunculata, which in no way dif- 
fers from the leaf variations of M. pertusa, matched by such Guate- 
malan collections as Steyermark 44832 and 45022, Standley 53214 and 
54465 from Honduras, Standley 9026 from Nicaragua, and Austin 
Smith Hi 75 from Costa Rica. The length of the peduncle in M. longi- 
pedunculata also falls within extremes of length occurring in M. pertusa. 
Some collections with subincised leaf blades have been misidentified 
as M. dilacerata, a South American species ranging northward to 
Panama. 

The decorticated part of the dried aerial roots of this species is 
commonly woven into hats, chair-seats, and other articles. The dried 
roots themselves are often used to make string or rope. 

Monstera tuberculata Lundell, Lloydia 2: 78. pi. 1. 1939. 

Epiphytic on trees in wet forest, 300 meters or less; Alta Verapaz; 
Izabal; Guatemala (?; perhaps in cultivation). Southern Mexico 
(Oaxaca; Veracruz; Tabasco); British Honduras, the type from Val- 
entin, El Cayo District, C. L. Lundell 6238. 

A small or large, glabrous vine, the stems sometimes pendent from the branches 
of trees and forming dense masses; petioles mostly about 3 cm. long, broadly winged 
for almost their whole length, the wings united below the base of the blade and 
forming a linear appendage 3.5-4.5 cm. long or shorter; leaf blades ovate or lance- 
ovate, thick, 10-16 cm. long, 5-9 cm. wide, acute or acuminate, shallowly cordate 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 335 

at the base and unequal, the nerves all very slender and equal, very numerous; 
peduncles slender, 5-6.5 cm. long; spathe about 8 cm. long, cuspidate-acuminate; 
spadix oblong, 5.5-8 cm. long, 2.5-3.5 cm. broad; pistils oblong-ovoid, narrowed 
into a conical style 4-5 mm. long, the spadix thus appearing densely tuberculate, 
the stigmas very small for the genus. 

Called "hoja de corazon" in Veracruz, and "trepepoyo" in Oa- 
xaca. The epidermis of the older branches is brown and separates 
in large flakes from the stems in drying. 



MONTRICHARDIA Crueger 
Reference: Engler, Pflanzenreich IV. 23C: 121. 1911. 

Large and coarse, terrestrial plants, the caudices erect, rather densely leafy, 
simple or sparsely branched; petioles vaginate to the middle or higher, the sheath 
persistent, clasping at the base, produced at the apex into a ligule; blades sagittate; 
peduncles shorter than the leaves, usually solitary, rarely geminate; spathes large, 
thick, convolute below, the limb open, deciduous after anthesis; spadix erect, 
slightly shorter than the spathe, the pistillate portion cylindric, densely many- 
flowered, the staminate portion contiguous to the pistillate and much longer, 
densely many-flowered; flowers unisexual, monoecious, naked; staminate flowers 
with 3-6 stamens, these distinct, obpyramidal-prismatic, contiguous, truncate at 
the apex, the filaments almost none; anthers subsessile, 2-celled, the cells oblong, 
dehiscent by short, apical slits; pistil subprismatic-obovoid, sessile, 1-celled; ovules 
1-2 in the cell, ascending, anatropous; fruit large, baccate, spongious, excavate at 
the apex, radiately costate, 1-seeded; seeds obovoid, the testa smooth and brown. 

The genus consists of two species, the other in Bahia, Brazil. 

Montrichardia arborescens (L.) Schott, Arac. Betreff. 1: 4. 
1854. Arum arborescens L. Sp. PI. ed. 2. 371. 1763. Masica. 

Occasional in tidal swamps of the North Coast, usually growing 
in shallow water; Izabal. British Honduras to Panama, Lesser An- 
tilles, Venezuela, and the Guianas. Figure 57. 

Caudex as much as 3 meters high but usually lower, 1.5-2 cm. thick or toward 
the base much thicker, the internodes about 1 cm. long, smooth or slightly aculeo- 
late; petioles 20-30 cm. long, the sheath extending above the middle; blades 20-30 
cm. long or often much larger, deeply sagittate, the basal lobes retrorse, triangular- 
lanceolate, acuminate, the anterior lobe triangular or ovate-triangular, short-cuspi- 
date or obtuse, the primary costal nerves 3-4 on each side, united to form an obscure 
collective nerve close to the margin; peduncles half as long as the spathe; spathe 
oblong-ovate, cuspidate, 10-13 cm. long, 6-7 cm. wide, white; spadix very thick 
and stout, the staminate portion 7 cm. long, the pistillate part one-third as long; 
berries 1-1.5 cm. in diameter. 

The plant is a characteristic one on muddy banks of the coastal 
swamps. The massive stems often are supported by prop roots. 




336 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 337 

The large white spathes are conspicuous and handsome, suggesting 
those of the cultivated calla (Zantedeschia) . 

PHILODENDRON Schott 
Reference: Engler & Krause, Pflanzenreich IV. 23Db. 1913. 

Scandent, often very large and climbing over tall trees, often emitting long 
aerial roots, the internodes mostly elongate, the young plants with long-vaginate 
leaves; sheaths often produced at the apex into a ligule, the petiole terete or cana- 
liculate; blades herbaceous or coriaceous (in the dried state), various in form, entire 
or often lobate or parted, the lateral nerves all parallel, equal or the primary ones 
slightly stouter; peduncles usually short, the spathe fleshy, white, green, or colored, 
the tube convolute, cylindric or ventricose, persistent after anthesis and surround- 
ing the spadix, its blade cymbiform and ovate, oblong, or lanceolate, usually erect, 
reconvolute after fecundation; spadix erect, about equaling the spathe, sessile or 
short-stipitate, the pistillate portion densely many-flowered, cylindric, fleshy in 
fruit, the staminate portion sterile below, fertile above, withering and recurved 
in fruit; flowers unisexual, naked, the staminate with 2-6 stamens, these obpyram- 
idal-prismatic, truncate at the apex, the connective of the anther thick, the anthers 
extrorse, oblong or linear, emarginate at the base; ovary of the pistillate flower 
ovoid or obovoid, 2-several-celled, the ovules 1 to several; stigma sessile, hemi- 
spheric or lobulate; fruits baccate, crowded, 1-several-seeded; seeds ovoid-oblong 
or ellipsoid, straight. 

One of the largest genera of the family, with about 225 species, 
widely distributed in tropical America. The species are rather numer- 
ous in Guatemala, and the plants often abundant, especially at lower 
elevations in wet forests. Most of the plants are large vines, and 
they are often highly ornamental because of their large and handsome 
leaves. Even the inflorescences of some species are showy and rather 
attractive. 

Leaf blades shallowly or deeply lobed or parted. 
Blades 3-parted. 

Ovules several in each cell of the ovary P. anisotomum 

Ovules solitary in the cells P. tripartitum 

Blades pinnatifid, with more than 3 lobes. 
Leaf blades triangular-sagittate in outline, bipinnately parted or bipinnatisect, 

most of the divisions lobed or parted P. Warscewiczii 

Leaf blades cordate-ovate in outline, pinnately parted, most of the divisions 
usually long and simple P. radiatum 



FIG. 57. Montrichardia arborescens. a, Spadix (X \4), 6, Portion of surface 
of staminate spadix (X 5). c, Apex of flowering shoot with leaves (X 14). d, 
Stamen, ventral view (X 10). e, Stamen, lateral view (X 10). /, Stamen, dorsal 
view (X 10). g, Portion of surface of pistillate spadix ( X 4). h, Pistillate flower, 
lateral view (X 6). i, Pistillate flower with longitudinal section through ovary 
(X 6). 



338 FIELDIANA: BOTANY, VOLUME 24 

Leaf blades entire. 

Blades obtuse or rounded at the base, or merely subcordate. 

Sheath ending 3-10 cm. below base of blade; blades shallowly cordate or sub- 
cordate at base P. Popenoei 

Sheath extending quite to base of blade or ending within 3.2 cm. of base of 
blade; blades attenuate to rounded at the base. 

Lateral pairs of most prominent primary nerves on lower leaf surface gen- 
erally 5-9 (rarely 10); sheath of petiole extending almost or quite to 
base of blade, its adnate portion on the petiole ending 1-6 mm. below 
junction with leaf blade; spathe 8-11 cm. long; spadix 8.5-10 cm. long. 

P. guttiferum 

Lateral pairs of most prominent primary nerves on lower leaf surface more 
numerous and conspicuous, usually 10-16; adnate portion of sheath of 
petiole ending usually 10-32 mm. below junction with leaf blade (rarely 
ending 5 mm. below junction); spathe usually 12-15 cm. long; spadix 

usually 11-15 cm. (rarely 8.5) long P. guatemalense 

Blades deeply cordate or sagittate at the base. 

Leaf blades sagittate, the basal lobes somewhat divaricate and narrowed at 

the apex . P. sagittifolium 

Leaf blades ovate-cordate or rounded-cordate, the basal lobes not divaricate, 

broadly rounded at the apex. 

Lateral nerves of the leaves above the basal ones usually 2-3 on each side; 
basal sinus of the leaf acute, very narrow; cells of ovary many-ovulate. 

P. hederaceum 
Lateral nerves of the leaves above the basal ones usually 5-7 on each side; 

basal sinus of the leaf broad and open; cells of ovary 1-3-ovulate. 
Pistil elongated, the style much longer and narrower than the ovary; cells 

of ovary 2-3-ovulate; peduncle 3.5-5 cm. long P. Hoffmannii 

Pistil only slightly longer than broad, the style about as broad as the 
ovary; cells of ovary 1-ovulate; peduncle 6-20 cm. long. .P. Smithii 

Philodendron anisotomum Schott, Oesterr. Bot. Zeitschr. 179. 
1858. P. affine Hemsl. Diag. PI. Mex. 37. 1878 (type from Barranco 
Hondo, Volcan de Fuego, Sacatepe"quez, Salviri). Cabeza de tow; 
Conti; Contin; Choboc (Coban, Quecchi). 

Moist or wet forest, 1500 meters or lower; Chiquimula; Jalapa; 
Jutiapa; Santa Rosa; Escuintla; Sacatepe"quez ; Suchitepe'quez ; Re- 
talhuleu; Quezaltenango; San Marcos. Southern Mexico. 

A small or large vine; petiole slender, terete, 15-40 cm. long or more, attenuate 
upward; leaf blades subcoriaceous, 3-parted almost to the broadly cordate base, the 
segments separated by rounded sinuses, unequal, the central one much wider than 
the lateral ones, the midlobe oblong-lanceolate, acuminate, 12-24 cm. long and 
4-10 cm. wide or larger, the lateral lobes oblique and somewhat incurved-falcate, 
the primary lateral nerves of the midlobe 4-5 on each side, ascending; peduncles 
stout, 5-8 cm. long or more; spathe green or bluish green outside, sometimes tinged 
with pink, dark red within, oblong, constricted below the middle, 6-12 cm. long; 
spadix pale yellowish, short-stipitate, cylindric, obtuse, 5-9 cm. long, 1-1.5 cm. 
thick; pistils cylindric, 4-6 mm. long, 5-6-celled, the cells 4-5-ovulate. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 339 

Called "anona conte" and "anona de conde" in El Salvador. 

This handsome plant is frequent at many places along the Pacific 
foothills and ascends rather high in the mountains where there is suf- 
ficient moisture. It is much planted for ornamental purposes in Gua- 
temala, especially as a pot plant in patios. The local name "cabeza 
de toro" ("bull-head") is an apt one, for the pendent leaf blades, with 
a broad central division and a narrow curved one on each side, do 
suggest the head and horns of a bull. 

Philodendron guatemalense Engler, Bot. Jahrb. 26: 514. 1899. 
P. belizense Standl. Field Mus. Bot. 11: 129. 1932 (type from base of 
Cockscomb Mountains, British Honduras, W. A. Schipp 545). P. 
Rojasianum Standl. & Steyerm. Field Mus. Bot. 23: 3. 1943 (type 
from Finca Pirineos, lower southern slopes of Volcan de Santa Maria, 
Quezaltenango, Steyermark 33244)- 

Epiphytic or growing upon rocks, usually in wet forest, ascending 
from 120 meters to about 1800 meters; Alta Verapaz; Izabal; Santa 
Rosa; Escuintla (type from Torola, J. D. Smith 2240) ; Sacatepe"quez; 
Chimaltenango; Solola; Suchitepe"quez; Retalhuleu; Quezaltenango; 
San Marcos. Chiapas; British Honduras. 

Usually a large vine, the internodes terete and stout; petiole 10-30 cm. long, 
with a conspicuous, broad, green sheath up to 1 cm. broad, this ending usually 
1-3.2 cm. below the base of the blade, sometimes produced at the apex into a broad 
green ligule as much as 2.5 cm. long; blades thin (when dried), oblong-ovate or 
elliptic-oblong, 15-38 cm. long and 8-17 cm. wide or even larger, obtuse or rounded 
at the cuspidate-acuminate apex, rounded or obtuse to very shallowly cordate at 
the base, the primary nerves 10-16 on each side, much stouter than the secondary 
ones, divergent at a wide angle; peduncles stout, 1.5-2 cm. long; spathe whitish, 
green, or yellowish green, about 12-15 cm. long and 2 cm. thick, the blade lanceo- 
late, attenuate; spadix green, slender, slightly attenuate upward, obtuse, short- 
stipitate, usually 11-15 (rarely 8.5) cm. long, shorter than the spathe; pistils 
3-5-celled, many-ovulate; stamens 2-3. 

A common and rather handsome plant, sometimes growing on 
rocks. Illustrated by Engler, Pflanzenreich IV. 23Db: /. 5. 1913. 
Called "anona conte" in El Salvador. 

In Guatemala this species ascends to 1800 meters on the lower 
slopes of some of the volcanoes, and in the mountains of Alta Vera- 
paz to elevations of 1600 meters. A related species, P. guttiferum, 
is found at lower elevations. 

The character of the prolonged free ligule of the sheath, used to 
distinguish P. Rojasianum, has been found to be a variable one. 
Steyermark 47918 from Volcan de Atitlan has the ligule well devel- 



340 FIELDIANA: BOTANY, VOLUME 24 

oped, as in P. Rojasianum, while other collections referred to P. gua- 
temalense have the ligule much less developed at the free apical end. 

Philodendron guttiferum Kunth, Enum. 3: 51. 1844. P. Tala- 
mancae Engler, Bot. Jahr. 26: 511. 1899 (type from Talamanca, Costa 
Rica, Pittier & Durand 9223}. P. calderense Krause in Engler & 
Krause, Pflanzenreich IV. 23Db: 8. 1913 (type from El Boquete, 
Panama, Pittier 3150}. P. linearipetiolatum Matuda, Madrono 10: 
49. fig. 1, a-b. 1949 (type from Finca Corcega, Chiapas, Mexico, 
Matuda 17781}. Conte. 

Epiphytic in wet forest, ascending from sea level to about 600 
meters; Alta Verapaz; Izabal; Huehuetenango. Southern Mexico 
(Oaxaca, Vera Cruz, Chiapas); Honduras; Nicaragua; Costa Rica; 
Panama; South America. 

A small or large, epiphytic vine, the branches stout, terete, with elongate or 
short internodes, the uppermost short, 8-10 mm. thick; petioles mostly 5-17 cm. 
long, varying from one-fourth the length of the blade to equaling the blade in 
length, the broad green sheath ending 1-6 mm. below the junction with the leaf 
blade or extending to the very base of the blade and conspicuously produced be- 
yond it as a ligule; leaf blades firm-membranaceous, thin, but often rather stiff, 
oblong, ovate-oblong, oblong-lanceolate, or rarely lanceolate, 10-20 cm. long and 
1.5-10 cm. wide, abruptly short-acuminate or cuspidate-acuminate, obtuse and 
somewhat oblique at the base, the primary lateral nerves 5-10 on each side, incon- 
spicuous but stouter than the secondary ones; peduncles terete, stout, mostly 2 cm. 
long or less to subsessile; spathe greenish- or creamy- white, ovoid- or elliptic- 
oblong, short-acuminate, convolute, 8-15 cm. long, 5 cm. wide, 2-3 cm. in diam- 
eter below; spadix short-stipitate or subsessile, the pistillate portion in anthesis 
about 4.5 cm. long, almost 2 cm. thick, in fruit 10 cm. long and 4 cm. thick, the 
staminate portion of equal length to twice as long, attenuate upward; pistil pris- 
matic, 3-4 mm. long; berries oblong, 6-7 mm. long, 4 mm. thick, 4-celled, the cells 
mostly 4-seriate, 2-3-seeded. 

This species is found in only a few departments of Guatemala, 
occurring at low elevations. Called "chupa-pito" in Chiapas. 

It has not been possible to separate P. guttiferum from P. calde- 
rense, P. Talamancae, and P. linearipetiolatum. In the Pflanzen- 
reich P. guttiferum is separated on the basis of its short petioles, half 
as long as the blades, but the illustration in that work (fig. 3, A-G) 
shows petioles scarcely if at all shorter, proportionately, than those 
of P. calderense. The extent to which the sheath reaches the base of 
the blade and the degree to which the lobes of the sheath are pro- 
duced beyond the insertion have been found to be variable charac- 
ters, as have also the relative proportions of the leaf blade and petiole 
length, and cannot be depended upon for specific segregation. It is 
probable that after an intensive monographic study has been made 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 341 

within this group, many more names considered as distinct species 
by Engler and Krause will have to be reduced to synonymy under 
P. guttiferum, P. lingulatum, and other species. Matuda himself 
(Las Araceas Mexicanas, Anal. Inst. Biol. Mex. 25: 146. 1954) re- 
duces his P. linearipetiolatum to synonymy under P. seguine Schott. 
The latter taxon Dr. Birdsey considers distinct from the form gener- 
ally treated as P. guttiferum. 

In describing P. linearipetiolatum, Matuda was misled by the 
winged nature of the sheath of the petiole and the relatively narrow 
leaf blades. The breadth and shape of the leaf blade vary on the 
same plant from juvenile to mature leaves. This can be verified by 
a study of preserved collections in the Chicago Natural History Mu- 
seum Herbarium. Such type of variation may be found in Steyer- 
mark 41771 from the Rio Tameja of the Department of Izabal, and 
Steyermark 41 733 from the Rio Bonita in the same region. The latter 
collection and Steyermark 49277 from the Rio Ixcan in the Depart- 
ment of Huehuetenango have blades at least as narrow as or narrower 
than those of the type collection of P. linearipetiolatum. Moreover, 
in deciding that the petiole in P. linearipetiolatum was winged in the 
natural state, while P. calderense was not, Matuda confused his col- 
lections of the latter species with P. guatemalense. He believed that 
the "sheathed petiole of P. calderense in its natural state is so com- 
pletely closed that it seems like a cylindrical tube, and this is the 
reason that the plant is called chupa-pito by the natives" (Madrono 
10: 50. 1949). However, the collections of Matuda (16697, 17707, 
and 17667) bearing the local name of "chupa-pito" are not P. calde- 
rense, but actually P. guatemalense. The manner of pressing and the 
rapidity of drying undoubtedly affect the appearance of the winged 
petiole, so that the wing appears more pronounced and better devel- 
oped when the petiole is pressed out flatly and evenly. The variation 
of the wingedness of the petiole on the same plant may be seen by 
referring to Steyermark 41771, discussed above, and the comparative 
narrowness of the wing is shown to good advantage in Steyermark 
41733. Although Matuda believed that the sheathed petiole was 
completely closed in the natural state of P. calderense (actually P. 
guatemalense), an examination of ample material from Guatemala of 
P. guatemalense reveals many collections with narrow, seemingly 
closed types of sheathed petioles as well as expanded broadly winged 
types (the latter to be noted in Steyermark 33244, the type collection 
of P. Rojasianum, Steyermark 37448, Steyermark 47918, Standley 
90818, and several others). 



342 FIELDIANA: BOTANY, VOLUME 24 

Philodendron hederaceum (Jacq.) Schott, Melet. 1: 19. 1832. 
Arum hederaceum Willd. Sp. PL 4: 486. 1805. P. cordatum Hort., 
not P. cordatum (Veil.) Kunth. P. scandens C. Koch & H. Sello, 
Ind. Sem. Hort. Berol. App. 14. 1853. P. oxycardium Schott, Syn. 
Aroid. 82. 1856. P. Miduhoi Matuda, Rev. Soc. Mex. Hist. Nat. 
11:95. 1950. 

Wet forest of the Pacific coast, at or little above sea level; Santa 
Rosa; Suchitepe'quez; Retalhuleu. Southern Mexico to West Indies 
and northern South America. Venezuela. 

A large or small vine, growing over trees or rocks, the internodes of the stems 
8-15 cm. long, 1 cm. thick or less; petiole semiterete, 10-18 cm. long or more; 
blades thick when dried, broadly cordate-ovate, mostly 20-30 cm. long and 16-20 
cm. wide but sometimes considerably larger, abruptly cuspidate, deeply cordate at 
the base, with a narrow acute sinus, the basal lobes broadly rounded, semiorbicular, 
the costa stout, with 3 basal and 2-3 costal nerves on each side, these much stouter 
than the secondary and tertiary nerves, divergent at a wide angle; peduncles 2-3 
cm. long or sometimes much longer, stout; spathes green or purplish, the tube 
5-6 cm. long, 3 cm. broad, the limb of about the same length; spadix cylindric, 
short-stipitate, the pistillate part 4-5 cm. long, 1.5 cm. thick, the staminate portion 
6-7 cm. long; pistils oblong, 5-6-celled. 

Illustrated by Engler, Pflanzenreich IV. 23Db:/. 19, A-G. 1913. 
The Maya name is reported from Yucatan as "acalcumche." The 
Guatemalan specimens are all sterile and their determination there- 
fore is uncertain. Some of the sterile material identified as P. Hoff- 
mannii may belong here. 

According to Dr. Birdsey, this is the common Philodendron sold 
in florists' shops and department stores everywhere, which long has 
passed under the name P. oxycardium and P. cordatum, although the 
latter name has been misapplied. 

Philodendron Hoffmannii Schott, Oesterr. Bot. Zeitschr. 178. 
1858. P. apocarpum Matuda, Madrono 10: 171. 1950 (type from 
Chiapas, Mexico) . Cola de coyote ( Jutiapa) ; Chile de chucho; Guaca- 
mayo; Quequesque de coyote. Figure 58. 

Epiphytic in moist or wet forest or sometimes on rocks or terres- 
trial, 800-2500 meters; Alta Verapaz; Zacapa; Chiquimula; Jutiapa; 
Jalapa; Santa Rosa; Escuintla; Guatemala; Chimaltenango; Re- 
talhuleu; Quezaltenango; San Marcos. San Luis Potosi to Yucatan 
and Chiapas; British Honduras; Honduras; El Salvador; Costa 
Rica; Panama. 

A small or large vine; internodes of the stem about 1 cm. thick; petioles sub- 
terete, 10-20 cm. long; leaf blades subcoriaceous, but thin when dried, ovate- 




FIG. 58. Philodendron Hoffmannii. a, Spadix (X %). b, Staminate flower 
viewed from above (X 12). c, Staminate flower (X 12). d, Part of flowering 
branch and leafy stem (X 1 A). e, Pistillate flower (X 

343 



344 FIELDIANA: BOTANY, VOLUME 24 

triangular, ovate-cordate or rounded-cordate, 12-24 cm. long, 10-20 cm. wide, 
cuspidate-acuminate, deeply cordate at the base, with a wide open sinus, the basal 
lobes rounded, the primary lateral nerves 3-6 on each side, divergent at a wide 
angle, much stouter than the secondary and tertiary ones; peduncles short or usu- 
ally elongate, 3.5-5 cm. long, stout; tube of the spathe ovoid, 3-5 cm. or more long, 
yellowish green outside, creamy white within, the limb ovate to oblong, cuspidate- 
acuminate, 4-15 cm. long; spadix sessile, the pistillate portion 1.5-6 cm. long, 2-4 
cm. thick, the staminate 4-10 cm. long and 2-2.5 cm. thick; pistils 4-12 mm. long, 
1-1.5 mm. thick, 3-4-celled, the ovules 2-3 in each cell, inserted near the base of 
the cell; stamens usually 4; berries ovoid, 7-8 mm. long, 3 mm. thick. 

Much of the Guatemalan material has been previously misidenti- 
fied as P. Pittieri Engler, which, according to present studies of Dr. 
Birdsey, is an anomalous species, not found in Guatemala. 

Philodendron apocarpum Matuda cannot be differentiated from 
P. Hoffmannii. Matuda used characters of longer ovaries, longer 
petioles and peduncles, fewer primary nerves of the leaf blades, and 
altitudinal differences in habitat in separating his species. These 
differences break down upon examination of numerous specimens, 
the more mature flowers, for example, showing longer ovaries. In 
fact, the character of an elongated pistil, narrowly contracted to the 
stigma, was originally used by Engler in setting aside a separate 
section, Macrogynium, for P. Hoffmannii. 

Called "anona conte" in El Salvador and "hoja de sereno" in 
Honduras. 

Philodendron lacerum (Jacq.) Schott has been reported doubtfully 
from Pete*n, but the report is probably based upon a misidentification 
of P. radiatum, according to Dr. Birdsey. P. Lundellii Bartlett has 
been listed as a Pete'n species, but the name is a nomen nudum, as no 
description has been published. 

Philodendron Popenoei Standl. & Steyerm. in Yuncker, Field 
Mus. Bot. 9: 264. 1940. 

Wet forest of the North Coast, at or little above sea level; Izabal 
(Entre Rios). Also on the Atlantic coast of Honduras, whence de- 
scribed. 

A large vine, climbing over trees, the internodes rather stout, elongate; petiole 
slender, 12-22 cm. long, narrowly winged by the sheath, this ending 3-10 cm. be- 
low the base of the blade; blades thin when dried, ovate or oblong-ovate, 16-33 cm. 
long, 13-20 cm. wide, short-apiculate, shallowly cordate or subcordate at the base, 
the basal lobes broadly rounded, short, the primary lateral nerves 8-9 on each side, 
slender, but conspicuously stouter than the others, the secondary and tertiary ones 
very numerous, equal, all divergent at a broad, often almost right, angle; peduncles 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 345 

stout, 3.5-4 cm. long; spathe green, 9-12 cm. long, 2 cm. thick, the blade broad, 
acuminate; spadix white, stipitate, the pistillate portion about 3 cm. long and 

1 cm. thick, the staminate part 5 cm. long; pistils columnar, scarcely more than 

2 mm. long. 

Called "hoja de sereno" in Honduras. 

Philodendron radiatum Schott, Oesterr. Bot. Wochenbl. 3: 
378. 1853. Memo de ledn; Uxb (Alia Verapaz). 

Wet forest, ascending to 2000 meters or perhaps even higher; Alta 
Verapaz; Izabal; Santa Rosa; Escuintla; Suchitepe"quez; Retalhuleu; 
Quezaltenango; San Marcos. Central and southern Mexico to 
Panama. 

A large or small vine, the stems thick, with short internodes; petiole pale green, 
subterete, 40-65 cm. long or more; blades thin-coriaceous when dried, the primary 
ones ovate-oblong, subentire, the next ones more or less incised or shallowly lobate, 
the adult blades pinnate-parted, ovate in outline, 35-50 cm. long and 30-35 cm. 
wide, or often much larger, the segments 5-10 on each side, linear-lanceolate, 
acuminate, the lower ones often more or less lobulate, the middle ones 2-3 cm. 
wide; peduncles stout, 4-6 cm. long; spathes cymbiform, bronze- or whitish-green, 
pale purple within, the tube ovoid, the blade oblong, apiculate, about equaling the 
tube; spadix 12-18 cm. long, the staminate portion slightly longer than the pistil- 
late, attenuate upward; pistils pale green, oblong, 2-3 mm. long, 8-10-celled, the 
cells chiefly 4-5-ovulate; stamens 4-6; berries oblong, 6-7 mm. long. 

Illustrated, Field Mus. Bot. 10: pi. 15. 1931; Pflanzenreich IV. 
23Db:/. 40. 1913. Called "hoja de peche" in El Salvador, "conte 
septina" in Chiapas. A striking plant because of the large and dis- 
tinctive leaves and densely draping tree trunks. It often is cultivated 
in patios of the central region, and it or a closely related species often 
is seen as a house plant in the United States. The leaves are much 
used in Guatemala as decorations on altars and elsewhere. The ju- 
venile leaves are very unlike the adult ones, being broadly cordate- 
ovate, the earliest ones nearly or quite entire, the succeeding ones 
undulate or very shallowly and obscurely lobate. 

Philodendron sagittifolium Liebm. Vid. Medd. Naturh. For. 
Kjoebenhavn 1850: 17. 1850. 

Epiphytic in moist or wet forest of the Occidente, 1200-1500 
meters; Quezaltenango; San Marcos. Southern Mexico. 

A large or small, coarse vine, the internodes of the stems more or less elongate, 
1-3 cm. thick; cataphylls linear-lanceolate; petioles slender or stout, often 40 cm. 
long or more, usually short- vagi nate; leaf blades oblong-sagittate, rather thick and 
when dry somewhat coriaceous, mostly 30-50 cm. long and 12-20 cm. wide, cuspi- 
date-acuminate, the basal lobes semioblong, separated by a deep, acute or obtuse 



346 FIELDIANA: BOTANY, VOLUME 24 

sinus, the basal nerves about 4 pairs, the costal nerves 4-5 pairs; peduncles 5-10 
cm. long or more, the spathe green or on the margins whitish, purplish within, as 
much as 17 cm. long and about 2.5 cm. thick; spadix sessile, the pistillate portion 
pale yellowish, 1 cm. thick, the staminate part whitish, 7-8 cm. long; pistils 2 mm. 
long, 5-7-celled; berries dirty yellow, obovoid, 3.5 mm. long. 

Philodendron Smithii Engler, Bot. Jahrb. 26: 540. 1899. 
P. Donnell-Smithii Engler ex Donn. Smith, Enum. PI. Guat. 2: 77. 
1891, nomen. Chuyuac, Okil (PeteX fide Lundell). 

Epiphytic in wet forest, ascending from sea level to about 1500 
meters; Pete*n; Alta Verapaz; Izabal (type from Rio Dulce, J. D. 
Smith 1535} ; reported from Quezaltenango. British Honduras; Hon- 
duras^); Nicaragua(?). 

A large vine, the upper internodes 3-4 cm. long; petioles terete, 45 cm. long; 
leaf blades coriaceous or subcoriaceous, subtriangular-cordate-ovate, about 40 cm. 
long and 28 cm. wide, short-acuminate, deeply cordate at the base, the sinus open, 
broad, oblong, the basal lobes rounded, about 10 cm. long and broad, the primary 
lateral nerves on each side about 4 basal and 3 costal ones, divergent at an angle 
of about 60 degrees; peduncles often elongate and 20 cm. long; tube of the spathe 
ovoid, 5-7 cm. long, 3 cm. broad, the limb ovate, short-cuspidate, 6-7 cm. long, 
5 cm. wide, or the spathe frequently larger, pale green outside, rose-red or brick-red 
within; spadix short-stipitate, the pistillate portion cylindric, 2.5-3.5 cm. long, 
10-12 mm. thick, the staminate portion cylindric, obtuse, 6-7 cm. long, 1 cm. 
thick; pistils ovoid, 1.5 mm. long, 8-celled, the cells 1-ovulate. 

The large leaves of this or some other species are much used in 
Coban for wrapping articles of food. 

Philodendron tripartitum (Jacq.) Schott, Melet. 1: 19. 1832. 
Arum tripartitum Jacq. Hort. Schoenbr. 2: 33. pi. 190. 1797. Contin. 

Wet forest, 1500 meters or lower; Alta Verapaz; Izabal; reported 
from Santa Rosa; San Marcos. Chiapas; British Honduras to Pan- 
ama; Jamaica; South America. 

A large or small vine, the internodes of the caudex 5-12 cm. long, 1-1.5 cm. 
thick; petioles terete, 20-30 cm. long, attenuate upward; blades thin-coriaceous 
when dried, 3-parted almost to the base, the divisions subequal, 15-25 cm. long, 
4-7 cm. wide, the midlobe oblong-lanceolate, cuspidate-acuminate; peduncles soli- 
tary, 3-5 cm. long; spathes whitish or pale green, yellowish above, the tube oblong, 
3-4 cm. long, the blade ovate or ovate-oblong, 5-6 cm. long, short-acuminate, 
about 3 cm. wide; spadix borne on a stipe 2-4 cm. long, the pistillate portion 3-4 
cm. long, 8-10 mm. thick, the staminate part 3.5-5 cm. long, slightly attenuate 
upward; pistils cylindric, 1.5-1.8 mm. long, 7-11-nerved, the cells 1-3-ovulate; 
berries red. 

Dr. Birdsey has suggested that the Guatemalan material may be 
P. Fenzlii Engler. However, as it has not been possible to place re- 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 347 

liance on the relative length of the lateral leaflets, a character used 
for separating P. Fenzlii from P. tripartitum, the Guatemalan mate- 
rial has been placed tentatively in P. tripartitum. 

Philodendron Warscewiczii C. Koch, Ind. Sem. Hort. Berol. 
App. 14. 1855; Engler, Pflanzenreich IV. 23Db:/. U. 1913. Guaca- 
mayo; Cupapayo; Mano de Ie6n. 

Usually in rather dry or sometimes moist or wet forest, generally 
growing on rocks but sometimes epiphytic, 1900 meters or lower; 
Zacapa; Jalapa; Jutiapa; Santa Rosa. El Salvador to Panama. 

A small or large vine, the caudex very stout, 7-10 cm. in diameter, the inter- 
nodes rather long; petiole subterete, 60-80 cm. long, 2 cm. thick; blades triangular- 
sagittate in outline, bipinnately parted or bipinnatisect, the principal lateral seg- 
ments 25-30 cm. long, the segments acute or attenuate, 1-2 cm. wide; peduncle 
4-5 cm. long; spathe scarcely constricted, purple within, 15 cm. long, 4 cm. in 
diameter, convolute; spadix sessile, the pistillate portion cylindric, 4 cm. long, 
almost 2 cm. thick at the base, the staminate part 10 cm. long, 1.5 cm. thick; 
pistils elongate-cylindric, 6-celled, 5 mm. long, 2 mm. thick; berries cylindric, 8 mm. 
long, the cells 3-4-seeded; seeds 2.5 mm. long. 

Known in El Salvador by the names "copapayo" and "ocopa- 
payo," and in Nicaragua as "papaya de monte." The large fruits 
are very sweet, juicy, and edible. When growing in dry, exposed 
places this is practically a deciduous plant, the large leaves turning 
yellow as the dry season advances, withering, and falling from the 
caudex. At the height of the verano the plants may be quite leafless. 

PISTIA L. Water Lettuce 
Reference: Engler, Pflanzenreich IV. 23F: 258. 1920. 

Floating, aquatic herbs, acaulescent, emitting numerous long roots; leaves spi- 
rally arranged in a rosette, spongy, broad, densely covered with minute, few-celled 
hairs; inflorescences very shortly pedunculate, the spathes foliaceous, whitish, gla- 
brous within, pilose outside, the pistillate inflorescence 1-flowered, the staminate 
2-8-flowered; flowers unisexual, naked; staminate flowers with 2 stamens, these 
short-connate, the anther cells opening by 2 vertical slits; ovary of the pistillate 
flower 1-celled, ovoid; ovules numerous, orthotropous, dense, 4-6-seriate; style 
short, terminal, the stigma obtuse, penicillate; fruit small, baccate, ovoid, with few 
or numerous seeds, irregularly dehiscent; seeds cylindric, subtruncate at the apex, 
excavate at the middle. 

A single species is known, widely distributed in tropical and sub- 
tropical regions of the earth. 

Pistia Stratiotes L. Sp. PI. 963. 1753. Lechuga;Lechuga de agua; 
Xicinchah (Pete"n, Maya, fide Lundell). Figure 59. 



348 



FIELDIANA: BOTANY, VOLUME 24 



Generally distributed in quiet fresh water, especially about the 
margins of lakes and ponds, 1200 meters or lower, or even occa- 
sionally at higher elevations; Pete*n; Izabal; Zacapa; Escuintla; 
Guatemala; Quezaltenango (in pond near Quezaltenango, perhaps 




FIG. 59. Pistia Stratiotes. a, Habit (X Yz). b, Flower (X 2). c, Flower 
with part of spathe removed ( X 2). d, Longitudinal section through ovary ( X 3). 



introduced). Ranging from southern Texas and Florida southward 
throughout all warmer parts of America; also in the Old World tropics. 

Leaves numerous, mostly 4.5-8 cm. long, very thick, soft, and spongy, pale 
green, cuneate-obovate, broadly rounded or emarginate at the apex, sessile. 

Known in El Salvador by the names "lechuga de sapo," "disci- 
plinilla," "repollo de agua," and "verdolaga de agua;" the Maya 
name is reported from Yucatan as "ibinha." The plant is common 
in all the lowland waters of Guatemala, as in Central America gen- 
erally. It is easily recognized by its rosette-like form and floating 
habit. In general appearance it has no resemblance to any other 
member of the Araceae. The inflorescences are so small that they 
are seldom seen or noticed. The plants often are grown in aquaria 
in the North. The most usual name for them in Central America 
is "lechuga de agua." 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 349 

RHODOSPATHA Poeppig 

Reference: Engler & Krause, Pflanzenreich IV. 23B: 90. 1908. 

Epiphytic vines, rooting at the nodes; leaves distichous, the petioles about 
equaling the blades, the sheath elongate, clasping at the base, the petiole geniculate 
and slightly thickened below the blade, the blades oblong-elliptic, thin, slightly 
inequilateral, the primary and secondary nerves numerous, parallel, spreading or 
ascending, arcuate near the margin; peduncles shorter than the leaves, the spathe 
broadly ovate or oblong-ovate, abruptly cuspidate, longitudinally nerved, green 
outside, whitish or pink within, soon deciduous; spadix stipitate, cylindric, densely 
many-flowered, sometimes with only pistillate flowers at the base, most of the 
flowers perfect; stamens 4, the filaments rather broad, complanate, narrowed to a 
slender, acuminate connective, the anthers rather broad, the cells elliptic, longer 
than the connective; fertile ovary quadrangular, 2-celled, the ovules several in 
each cell; stigma linear or rarely 2-3-lobate; berries small, cylindric-prismatic, 
truncate, 2-celled, 10-12-seeded; seeds vertically imbricate, rounded-reniform, 
lenticular. 

About 10 species are known, distributed from Guatemala to Bra- 
zil. One other Central American one is recorded for Costa Rica. 

Sheath of the petiole extending upward to the node of the petiole R. nervosa 

Sheath of the petiole ending some distance below the node R. Tuerckheimii 

Rhodospatha nervosa Lundell, Contr. Univ. Mich. Herb. 6: 
3. 1941. 

Wet forest, about 150 meters; British Honduras (type from Mid- 
dlesex, Stann Creek District, P. H. Gentle 2797; also Cockscomb 
Mountains, Schipp Si 26). 

A large epiphytic vine; petiole broadly vaginate below, the sheath extending 
to the base of the node, the node about 2.5 cm. long; leaf blades stiff-membrana- 
ceous, ovate-oblong or broadly ovate, 45-60 cm. long, 20-35 cm. wide, obtuse or 
rounded at the apex and short-cuspidate, broadly rounded at the base, sometimes 
abruptly short-decurrent, the primary lateral nerves about 40 on each side, very 
slender, prominent beneath; peduncles stout, 8-15 cm. long; spathe deciduous; 
stipe of the spadix 1 cm. long; spadix at anthesis 11 cm. long, 11 mm. in diameter, 
in fruit about 16 cm. long and 1.5 cm. in diameter; stamens 3-4.5 mm. long, the 
anthers 1.2 mm. long; ovary 2 mm. long, the ovules numerous; style about equal- 
ing the ovary, the stigma elevated. 

Rhodospatha Tuerckheimii Engler & Krause, Pflanzenreich 
IV. 23B:92. 1908. 

Wet, mixed forest, 1200 meters or lower; Alta Verapaz (type col- 
lected at 1200 meters, Tuerckheim 1103); Izabal (Entre Rios and 
Montana del Mico). Atlantic coast of Honduras. 



350 FIELDIANA: BOTANY, VOLUME 24 

A large epiphytic vine, climbing over trees; petioles about 20 cm. long, the 
sheath coriaceous, persistent, ending about 3 cm. below the base of the blade; leaf 
blades 25-40 cm. long, 9-17 cm. wide, abruptly short-cuspidate, obtuse or acute at 
the base, the primary nerves 12-14 on each side, divergent at a wide angle; pedun- 
cles slender, about 25 cm. long; spathe white, oblong, short-acuminate, 10 cm. long, 
3.2 cm. wide; spadix cylindric, obtuse, slightly attenuate upward, 9 cm. long, 1 cm. 
thick, pale purplish; pistils 4 mm. long, 1.5 mm. wide, the apex hexagonal, trun- 
cate, crowned by the small rounded stigma. 



SPATHIPHYLLUM Schott 
Reference: Engler & Krause, Pflanzenreich IV. 23B: 118. 1908. 

Plants terrestrial, acaulescent; leaves equitant, thin, the petioles long and 
slender, often geniculate at the apex, vaginate to the middle or higher, the blade 
oblong to elliptic, cuspidate-acuminate, the costa stout, the primary and secondary 
nerves numerous, parallel, slender, spreading or ascending, arcuate near the mar- 
gin; peduncles as long as the leaves or longer, the spathe cuspidate, decurrent upon 
the peduncle, membranaceous, at first convolute but soon explanate, withering 
and persistent; spadix sessile or stipitate, cylindric, erect, densely many-flowered, 
shorter than the spathe, flowering from below upward; flowers perfect, 3-parted 
or rarely 2- or 4-parted; sepals fornicate above and subtruncate, coherent or con- 
nate to form a truncate cup; stamens as many as the sepals and opposite them, the 
filaments short, dilated toward the apex, the anthers ovoid, the cells oblong, longer 
than the connective; ovary oblong, usually 3-celled, the ovules in each cell 2-8; 
style continuous with the ovary, conically elongate and protruding beyond the 
perianth, or almost obsolete; stigma 3-4-lobate, sessile; fruit baccate, rounded at 
the apex or conic-attenuate, 3-celled, the cells 1-8-seeded; seeds oblong, somewhat 
curved and more or less reniform. 

About 25 species are known, ranging from southern Mexico to 
Brazil, with one species in the Philippines. A few others are known 
from Central America. The young inflorescences are much eaten in 
some parts of the Pacific coast of Central America, usually fried 
with eggs. 

Cells of the ovary 2-ovulate; sheath ending well below the node of the petiole. 

S. phryniifolium 

Cells of the ovary 4-8-ovulate; sheath extending almost or quite to the node of the 
petiole S. blandum 

Spathiphyllum blandum Schott, Oesterr. Bot. Wochenbl. 7: 
159. 1857. Huisnay; Guisnay. Figure 60. 



FIG. 60. Spathiphyllum blandum. a, Pistil with sepals (X 43^). 6, Stamen 
(X 4K)- c > Ovule (X 52). d, Leaf blade and portion of sheath attached to 
petiole (X 1 A). e, Lower portion of plant (X YZ). f, Spadix with most of spathe 
removed (X 1). 




351 



352 FIELDIANA: BOTANY, VOLUME 24 

Wet forest, 1500 meters or lower, most frequent at low elevations; 
type from Guatemala, Friedrichsthal, the locality unknown; Alta 
Verapaz; Izabal; Huehuetenango. British Honduras; Honduras. 
Southward to Colombia. 

Plants about a meter high; petioles 20-40 cm. long, the apical node 2-2.5 cm. 
long, the sheath broad, reaching almost or quite to the node; blades mostly oblong- 
elliptic, 20-35 cm. long, 10-20 cm. wide, abruptly short-cuspidate or acute or short- 
acuminate, acutely contracted at the base; peduncles 30-50 cm. long or more, very 
slender, the spathe pale green or yellowish, elliptic or obovate, 15-20 cm. long, 
6-7 cm. wide, caudate-acuminate, long-decurrent upon the peduncle; spadix whit- 
ish, cylindric, obtuse, 5-7 cm. long, at maturity as much as 2 cm. thick; pistils 
ovoid-conic, 6 mm. long, the cells 6-8-ovulate; style conic, slightly exceeding 
the ovary. 

This species has been confused in Guatemala and elsewhere with 
S. Friedrichsthalii and S. Wendlandii, both of which are considered 
distinct by Dr. George S. Bunting, recent monographer of the genus. 

Spathiphyllum phryniifolium Schott, Oesterr. Bot. Wochenbl. 
7: 159. 1857. S. Ortgiesii Regel, Gartenflora 21: pi. 738. 1872. Gus- 
nay; Bushnay; Burnay; Guisnay; Huisnay. 

Moist or wet forest, of the Pacific boca costa, chiefly at low eleva- 
tions but ascending to 1400 meters in Quezaltenango; Santa Rosa; 
Escuintla; Suchitepe"quez; Quezaltenango; San Marcos. Chiapas 
(whence the type). 

Plants about a meter high, with few leaves; petioles about 40 cm. long, very 
slender, the portion above the node 3 cm. long, the sheath narrow, ending at some 
distance below the node, the blade broadly oblong or oblong-elliptic, 35-55 cm. 
long, 16-23 cm. wide, cuspidate-acuminate, contracted at the base and sometimes 
rather long-decurrent, the primary nerves numerous, ascending at an angle of 
about 70 degrees; peduncles slender, 60 cm. long or more, the spathe oblong-ellip- 
tic, about 15 cm. long and 5-6 cm. wide, green, long-decurrent on the peduncle, 
cuspidate-acuminate; spadix cylindric, rounded at the apex, 6.5-10 cm. long, 1.2- 
1.5 cm. thick, or in fruit much thicker; pistils 4-5 mm. long, the style long-produced 
above the perianth. 

The names "gusnay" and "huisnay" presumably are variants 
of a single name, of Nahuatl origin. The young inflorescences are 
often offered in the markets at Quezaltenango and elsewhere, but 
they are seldom served on the table in the hotels of Guatemala, at 
least in those frequented by tourists. They are gathered after the 
spathes have opened, and are tied in bunches for sale. An amusing 
tale is told in connection with this plant. Some few years ago a 
North American visiting Guatemala was served in some hotel with 
soup in which he discovered what he took to be diminutive ears of 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 353 

corn. Upon his return to the United States he reported the matter 
to the United States Department of Agriculture. Because of the 
hope of some day discovering the wild ancestor of corn, the Depart- 
ment was interested and sent two men to Guatemala to investigate 
the plant. Investigation revealed that the supposed corn consisted 
of the spadices of Spathiphyllum, which bear a remote resemblance 
to much-reduced ears of maize. 

STENOSPERMATION Schott 
Reference: Engler & Krause, Pflanzenreich IV. 23B: 81. 1908. 

Plants epiphytic, usually with elongate and often scandent caudices; petioles 
long-vaginate, nodose shortly below the apex; blades oblong-elliptic to lanceolate, 
asymmetric, the primary lateral nerves numerous, ascending; peduncles rather 
long, at first nutant at the apex, later erect; spathe convolute, finally open, navi- 
culiform, whitish, soon deciduous; spadix stipitate, cylindric, whitish; flowers all 
perfect, naked; stamens 4, the filaments complanate, abruptly narrowed at the 
apex into the slender connective, equaling the ovary, the anthers 2-celled, the cells 
oblong-ovoid, acute, laterally dehiscent by a longitudinal slit; ovary obpyramidal 
or prismatic, truncate at the apex, 2-celled; ovules 4 or more in each cell, ana- 
tropous; style short, thicker than the ovary, the stigma linear-oblong; fruit small, 
baccate, obovoid, subtruncate at the apex, 2-celled; seeds clavate-cylindric. 

The genus contains about 20 species, of which six are recorded 
for Central America. 

Leaf blades 30 cm. long or more; sheath of the petiole extending to the base of the 
blade S. multiovulatum 

Leaf blades mostly 30 cm. long or shorter; sheath of the petiole ending below the 
base of the blade S. robustum 

Stenospermation multiovulatum (Engler) N. E. Brown, 
Gard. Chron. III. 15: 684. 1894. S. Spruceanum var. multiovulatum 
Engler, Hot. Jahrb. 4: 281. 1885. 

Reported by Engler and Krause from rain forest above Purulha, 
Baja Verapaz, 1700 meters, F. C. Lehmann 1359. Colombia. 

Caudex stout, as much as 2 meters long; petioles 20-25 cm. long, vaginate to 
the apex; leaf blades rigid-coriaceous, oblong or narrowly elliptic-oblong, narrowed 
to the mucronate apex, cuneately contracted to the base, 30 cm. long or more, 10- 
14 cm. wide; peduncles 30-45 cm. long; spathe whitish, broadly elliptic, abruptly 
acuminate, 12-16 cm. long; spadix whitish, short-stipitate, 8-18 cm. long, 12-15 
mm. thick. 

Stenospermation robustum Engler, Bot. Jahrb. 37: 113. 1905. 

On trees or rocks, usually in wet forest, 200-1000 meters; Alta 
Verapaz; Huehuetenango. Costa Rica. 



354 FIELDIANA: BOTANY, VOLUME 24 

Caudex as much as 2 cm. thick, the internodes short; petioles slightly or much 
shorter than the blades, the sheath ending 0.5-4 cm. below the base of the blade; 
leaf blades subcoriaceous, oblong-elliptic, 15-30 cm. long and 5-10 cm. wide, or 
sometimes larger, short-acuminate, acute at the base; peduncles 30 cm. long; spathe 
almost pure white; spadix borne on a stipe up to 1 cm. long, 10 cm. long, 1.8 cm. 
thick; berries ovoid, many-seeded; seeds reniform. 

Illustrated, Pflanzenreich IV. 23B:/. 35, A-H. 1905. 



SYNGONIUM Schott 
Reference: Engler, Pflanzenreich IV. 23E: 121. 1920. 

Scandent epiphytes; leaves very variable in form according to the stage of de- 
velopment, the earliest blades ovate, the next sagittate, the adult blades trisect to 
pedatisect; petioles long, terete above, long-vaginate; peduncles equaling or shorter 
than the spathes, dependent in fruit; spathes usually yellowish or whitish green or 
purplish, the tube more or less ovoid, slightly longer than the pistillate inflores- 
cence, the blade oblong-ovate, open in anthesis; spadix much shorter than the 
spathe, the pistillate portion oblong-conoid, the staminate portion clavate, longer 
than the pistillate; flowers unisexual, naked; staminate flowers with 3-4 stamens, 
these connate to form a truncate-obpyramidal, 3-4-sided synandrium, the anthers 
linear, with a thick connective, dehiscent by a short slit below the apex of the con- 
nective; pistillate flowers connate, the ovary obovoid, 2-celled or by abortion 
1-celled; ovule solitary in the cell, erect, anatropous; stigma convex or hemi- 
spheric; fruits baccate, connate to form an ovoid syncarp; seeds obovoid. 

Engler recognizes fourteen species, six or seven of which are 
Central American. 

The following key to the genus was provided by Dr. M. R. 
Birdsey: 

Adult leaf blades of no more than three leaflets (laterals may have auricle) ; interior 

of spathe blade clear milky white; stigma bilabiate S. Salvador ense 

Adult leaf blades of more than three leaflets (intermediate stages of three leaflets 

may be found); interior of spathe blade whitish green; stigma discoidal. 
Anthers of male flowers completely fused into a "tooth-like" synandrium with 
a retuse apex; juvenile leaf blades variously marked with grayish white. 

S. angustatum 

Anthers of male flowers fused to varying degrees into a synandrium with a trun- 
cate apex; juvenile leaf blades very rarely variegated, if variegated then 
with a yellowish shade. 

Plant larger and thicker in all parts; adult leaf blades in cross section approxi- 
mately 325 microns thick; median leaflet 22-47 cm. long; juvenile leaves 

thick, cordate S. macrophyllum 

Plant smaller and thinner in all parts; adult leaf blades in cross section approx- 
imately 160 microns thick; median leaflet 17-38 cm. long; juvenile leaves 
thin, sagittate or hastate S. podophyllum 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 355 

Syngonium angustatum Schott, Oesterr. Bot. Zeitschr. 8: 178. 
1858. S. albolineatum Hort. in Engl. Arac. exs. et illustr. no. 243 
(circa 1883). S. podophyllum Schott, var. albolineatum (Hort.) 
Engler, Pflanzenreich IV. 23E: 129. 1920. 

Wet forest, epiphytic, near sea level ; Pete"n ; Izabal (near Puerto 
Barrios). Mexico (Tabasco); Costa Rica. 

Juvenile shoots: stems not glaucous, but with numerous, small, rough emer- 
gences, internodes very short, scarcely 1 cm. long in non-scandent stage, up to 
15 cm. or more long in scandent stage; petioles 3-22.5 cm. long, vaginate less than 
one-third the length; lamina variously marked with gray-green, the first-formed 
laminae cordate, later ones sagittate to hastate. Adult shoots: internodes very 
short in rosette stage, up to 8 cm. long and 1.5 cm. in diameter in later scandent 
stage, with numerous rough emergences; petioles vaginate from more than one- 
half to four-fifths their length; laminae dark green, occasionally with gray-green 
on midrib, trisect to 11-pedatisect, ultimate leaflet earlike; leaflets gradually, not 
abruptly, diminishing in size from central to lateral leaflets, elliptic to oblong to 
lanceolate; peduncle 7.8-13 cm. long, glaucous, tri-cornered ; spathe tube 2-4 cm. 
long, 1.2-2.6 cm. wide, elliptic, green within and without, persisting in fruit and 
becoming red to reddish orange; spathe blade 5.6-9 cm. long, 1.5-3 cm. wide, 
cream-colored within and without, deciduous after flowering of staminate part of 
inflorescence; pistillate inflorescence 1.3-2.7 cm. long, 0.4-1.1 cm. wide, dull green, 
oblique at base, becoming gradually attenuate; staminate inflorescence oblong, 
gradually tapering at base, obtuse at apex, 2.6-6.5 cm. long, 0.5-1.5 cm. wide, 
light yellow; pistillate flowers mostly irregularly hexagonal, stigma discoidal; stam- 
inate flowers completely fused into a synandrium, usually of 4 anthers, rarely 5, 
the apex markedly retuse, forming a tooth-like structure; sterile staminate flowers 
on the lower part of the inflorescence, more irregular and less retuse; fruit brown 
with darker brown flecks, 2-4.4 cm. long, 1.5-2.4 cm. wide; seeds dark gray. 

Dr. Birdsey has supplied the description of this species. 

Syngonium macrophyllum Engler, Pflanzenreich IV. 23E: 
128. 1920. 

Wet forest, epiphytic, mostly near sea level to 300 meters, rarely 
to 1000 meters; Izabal; Alta Verapaz. Mexico (Tabasco); El Sal- 
vador; Honduras; Costa Rica; Panama. Plants from Colombia may 
belong here. 

Juvenile shoots: stems glaucous, the internodes 0.8-5.2 cm. long, usually scan- 
dent; petioles 7.5-25 cm. long, glaucous, vaginate one-third to one-half the length; 
laminae coriaceous, ovate-cordate to ovate-sagittate to elliptic-hastate in more 
mature shoots. Mature shoots: stems scandent, glaucous, internodes 2.5-4 cm. 
long, up to 4 cm. thick; petioles glaucous, 25-78.5 cm. long, vaginate one-half to 
two-thirds the length; laminae coriaceous, medium green, pedatisect up to 9 leaf- 
lets; peduncle 10-20.5 cm. long, up to 8 inflorescences per leaf axil; spathe tube 
ovate, green within and without, glaucous, 3-5 cm. long, 1.5-2.2 cm. wide; spathe 
blade at first green, later light cream with greenish suffusion, apparently early de- 



356 FIELDIANA: BOTANY, VOLUME 24 

ciduous, oblong, short mucronate at apex, 6.7-9 cm. long, 1.4-2.4 cm. wide; pistil- 
late inflorescence gradually attenuate above, light green, 1.1-2 cm. wide (when 
fresh); staminate inflorescence oblong, cylindrical, gradually attenuate above, ob- 
tuse at apex, abruptly constricted at base just above the sterile staminate portion; 
sterile staminate portion about the width of the pistillate section, light brownish- 
yellow, 5.3-9 cm. long, 1.1-1.6 cm. wide (when dry), 6.8-10.4 cm. long, 1.5-2 cm. 
wide (when fresh); pistillate flowers irregularly pentagonal or hexagonal, stigma 
subsessile, discoid, yellowish at first, becoming black; staminate flowers of 4 an- 
thers fused into a synandrium, the line of fusion very slightly or not at all visible, 
apex truncate or retuse; sterile staminate flowers somewhat larger, the sides straight 
and not indented as in the fertile staminate ones; fruit large, 6 cm. long, 4.5 cm. 
wide, brown, with darker brown flecks; tube of spathe surrounding fruit at 
maturity, green within and without; seeds dark gray, 1.4-1.8 cm. long, 1.4 cm. 
wide. 

Plants of this species have frequently been confused with S. podo- 
phyllum, according to Dr. Birdsey, who has kindly supplied the 
description. 

Syngonium podophyllum Schott, Syn. Aroid. 68. 1856. 

Moist or wet forest, chiefly in the tierra caliente, ascending to 1350 
meters but most plentiful at lower elevations; Pete"n; Alta Verapaz; 
Izabal; Chiquimula; Jutiapa; Escuintla; Guatemala; Sacatepe*quez; 
Solola; Chimaltenango; Suchitepe"quez; Retalhuleu; Quezaltenango; 
San Marcos. Southern Mexico; British Honduras to El Salvador 
and Panama. 

A large or small epiphytic vine, the stems 1-1.5 cm. thick, the internodes up 
to 10 cm. long; petioles 20-50 cm. long, the sheath as much as 20 cm. long; leaf 
blades 5-11-cleft, the outer segments gradually smaller, the middle one about 20 
cm. long and 6-7 cm. wide, or often smaller, the outer ones oblong-lanceolate, ob- 
tuse to long-acuminate, distant, cuneate at the base; peduncles usually several 
together, about 10 cm. long; tube of the spathe 3-5 cm. long and 1.5-2 cm. thick, 
oblong-ovoid, green outside, the blade about 6 cm. long and 4-5 cm. wide, mostly 
whitish green or white or pink within, ovate, cuspidulate; pistillate inflorescence 
2 cm. long, green, the staminate 5 cm. long and 10-13 mm. thick, slightly atten- 
uate to each end; ovaries depressed-obpyramidal; fruiting spathes usually turning 
bright red. 

Called "pico de guara" in El Salvador. The Maya name "ochil" 
is reported from Yucatan. The plant is abundant in many localities, 
often growing on isolated trees in rather dry regions, for it withstands 
dry air better than most plants of the family. Perhaps on this ac- 
count it has proved successful as a pot plant in the United States; 
it often is planted out of doors in the southern United States. The 
large and abundant fruiting spathes are showy and rather hand- 
some. 



STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 357 

Syngonium salvadorense Schott, Oesterr. Bot. Zeitschr. 8: 
178. 1858. S. Donnell-Smithii Engler, Bot. Jahrb. 37: 141. 1905. 
Conte; Huevo de burro. 

Moist or wet forest, 1000 meters or less; Izabal; Chiquimula; 
Santa Rosa; Escuintla (type from Escuintla, J. D. Smith 2238}', 
Guatemala; Suchitepe"quez; Retalhuleu; Quezaltenango; San Mar- 
cos. Chiapas; El Salvador. 

A large epiphytic vine, the stems 1 cm. thick, the internodes 4-5 cm. long or 
more; petioles 20-30 cm. long, vaginate to the middle or higher; adult leaf blades 
trisect, the segments coherent at the very base, the middle segment ovate or 
broadly ovate, 15-20 cm. long, about 13 cm. wide, obtuse to acuminate, the lateral 
segments obliquely oblong or elliptic, slightly shorter than the middle one and 
about half as wide; peduncles 12 cm. long or less; tube of the spathe 5-6 cm. long, 
2-3.5 cm. broad, the limb ovate-oblong, acute, 5-7 cm. long, clear milky white 
within; pistillate spadix 2 cm. long, the staminate sterile portion 4.5 cm. long and 
above the middle almost 2 cm. thick; syncarp 5-6 cm. long and as much as 5 cm. 
thick; stigma bilabiate; seeds ovoid, 5-6 mm. long. 

In El Salvador called "anona conde," "anona conte," and "pico 
de guara." The spathes are sometimes bright red at maturity. The 
spadices are said to be edible when thoroughly ripe. 



UROSPATHA Schott 
Reference: Engler, Pflanzenreich IV. 23C: 30. 1911. 

Terrestrial herbs, growing in swamps, arising from horizontal or perpendicular 
rhizomes; leaves basal, few, the petioles very long, vaginate only at the base, the 
blades sagittate; peduncle terminal, the spathe erect, usually colored, convolute 
below, open above and gradually narrowed to the apex, persistent; spadix short- 
stipitate or sessile, several times shorter than the spathe, densely many-flowered; 
flowers perfect, perigoniate; sepals 4-6, fornicate, subtruncate at the apex; stamens 
4-6, the filaments rather broad, subcompressed, abruptly narrowed into the con- 
nective, scarcely longer than the ovary, the anthers exceeding the connective, the 
cells ovate-elliptic, dehiscent by an extrorse, apical slit; pistil truncate-conoid, in- 
completely 2-celled, the ovules 2 or more in each cell; berry surrounded by the 
accrescent perianth, 2-celled, or one of the cells aborted, 1-2-seeded. 

About 12 species are known, all American and chiefly Brazilian. 
Another Central American one occurs in Costa Rica. 

Urospatha Tuerckheimii Engler, Bot. Jahrb. 37: 121. 1905. 

Type from Livingston, Izabal, Tuerckheim I I.I 131; represented 
in the Herbarium of Chicago Natural History Museum by a photo- 
graph of the type (Negative no. 12195). Frequent in Manicaria 
swamps of the north coast (Izabal). Endemic. 



358 FIELDIANA: BOTANY, VOLUME 24 

Plants 1-1.5 meters high; petioles greatly elongate, vaginate for 20 cm.; blades 
sagittate, 60 cm. long or more, acuminate, the basal lobes equaling or shorter than 
the midlobe, somewhat divergent, separated by a broad sinus, more or less acumi- 
nate; peduncles about 50 cm. long; spathe bronze outside, yellow-green within, 
linear-lanceolate, twisted above, 20-35 cm. long, 3.5-6 cm. wide, narrowly long- 
attenuate, open almost to the base; spadix short-stipitate, 4-7 cm. long, 1.5-2 cm. 
thick, very obtuse, purplish green; sepals 4; cells of the ovary 2-ovulate. 

A rather showy but not especially handsome plant, its huge leaves 
succulent, soft and flabby. The plant in gross aspect is quite similar 
to U. sagittifolia, illustrated by Engler, Pflanzenreich IV. 23C:/. 12. 
1911. 

XANTHOSOMA Schott 

Reference: Engler, Pflanzenreich IV. 23E: 41. 1920. 

Terrestrial plants with thick rhizomes or tubers, sometimes with elongate cau- 
dices, the sap more or less milky; petioles long and thick, vaginate below, the blades 
sagittate or hastate, or pedately 3-many-cleft; peduncles solitary or several, usually 
short; tube of the spathe ovoid or oblong, convolute, persistent, constricted above, 
the blade oblong-cymbiform or oblong-lanceolate; spadix shorter than the spathe, 
the pistillate portion cylindric, narrowed upward, densely many-flowered, the 
sterile staminate portion longer than the pistillate, the fertile staminate part thick- 
cylindric, somewhat narrowed at the apex, twice as long as the pistillate or longer; 
flowers unisexual, naked; staminate flower with 4-6 stamens, these connate to form 
a truncate-obpyramidal synandrium, plane at the apex and somewhat 5-6-angu- 
late, the anther cells oblong or oblong-triangular; ovary ovoid, 2-4-celled, the styles 
thick, coherent as a ring; ovules several in each cell, anatropous, biseriate, erect; 
stigma discoid or hemispheric-discoid, 3-4-lobate; berries cylindroid, 3-4-celled, 
crowned by the impressed stigma, the cells many-seeded; seeds ovoid, the testa 
sulcate, the endosperm copious. 

About 40 species are known, extending over most of tropical 
America, chiefly at low elevations. Seven species are known from 
Central America. 

Leaf blades cleft into 3 or more segments. 

Outer lateral segments of the leaves with a rounded lobe at the base . .X. pedalum 

Outer lateral segments of the leaves not lobate at the base X. Hoffmannii 

Leaf blades sagittate, entire. 

Plants without tubers, the caudex developed above the soil, often greatly elon- 
gate, thick, and procumbent X. robustum 

Plants with a hypogaean, more or less tuberous caudex. 

Leaves glabrous X. violaceum 

Leaves more or less pubescent beneath X. mexicanum 

Xanthosoma Hoffmannii Schott, Oesterr. Bot. Zeitschr. 15: 
33. 1865. Xanthosoma Wendlandii Schott, Oesterr. Bot. Zeitschr. 
15: 33. 1865, nomen. Acontias Wendlandii Schott, Oesterr. Bot. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 359 

Zeitschr. 8: 178. 1858. A. Hoffmannii Schott, Prodr. Aroid. 196. 
1860. X. Hoffmannii var. Wendlandii (Schott) Engl. Fl. Bras. 3, 
pt. 2: 191. 1878. Quequescamote de culebra. 

Moist or wet, mixed forest, 700-1700 meters; Alta Verapaz; 
Chiquimula; Guatemala; Santa Rosa; Suchitepe"quez; Huehuete- 
nango. Southern Mexico; Honduras; Nicaragua; Costa Rica. 

Plants arising from a small tuberous hypogaean caudex; leaves glabrous, the 
petioles 20-30 cm. long; leaf blades reniform in outline, about 25 cm. long and 
30 cm. wide, pedatisect, the segments 5-7, the middle one oblong-elliptic, cuspi- 
date, arcuate-cuneate at the base, 16-20 cm. long, 7-10 cm. wide, the lateral seg- 
ments oblong-elliptic, the outermost smaller; peduncles about 10 cm. long, thicker 
than the petioles; tube of the spathe oblong, 5-6 cm. long, the limb oblong, cuspi- 
date-acuminate, white, almost 15 cm. long, 4.5-5 cm. wide; stipe of the spadix 
1 cm. long, the pistillate portion of the spadix 2.5 cm. long, the sterile staminate 
part 3-4 cm. long, the fertile staminate portion 7.5-10 cm. long; berries oblong, 
yellow, 1 cm. long, 4 mm. thick. 

Called "comida de culebra" in Costa Rica. 

Xanthosoma mexicanum Liebm. Vid. Medd. Kjoebenhavn 
15. 1850. Quequeshque de culebra. 

Wet forest, 1200-1800 meters; Huehuetenango (Rio Trapichillo, 
between La Libertad and Paso del Boqueron, Steyermark 51115). 
Southern Mexico; Nicaragua; Panama. 

Plants rather small and slender, arising from a small tuberous hypogaean cau- 
dex; petioles long and usually slender, puberulent or short- villous; leaf blades green 
and glabrous above, sparsely puberulent or short-villous beneath, broadly sagit- 
tate-triangular or rounded-ovate, commonly 20-30 cm. long and 15-25 cm. wide 
near the base, shortly cuspidate-acuminate, the basal lobes somewhat triangular 
or rounded, often narrowed to an obtuse apex, separated by a very broad and open, 
usually shallow sinus; peduncles 15-35 cm. long, sparsely pilose; tube of the spathe 
green and purple, about 6 cm. long and 2 cm. broad, oblong, somewhat pubescent, 
the limb lanceolate or ovate, acuminate, white, pubescent outside, 8-12 cm. long, 
4-5 cm. wide, cuspidate-acuminate; spadix somewhat shorter than the spathe, the 
staminate portion slender, about 8 cm. long. 

There is a popular belief in Huehuetenango that the fruits are 
poisonous to snakes. The fruits are put in places known to be fre- 
quented by snakes, and if the animals eat them, they die a very 
proper fate for serpents so foolish. 

Xanthosoma pedatum Hemsl. Biol. Centr. Amer. Bot. 3: 418. 
1885. 

Known only from the type, Sacatepe"quez, Barranco Hondo, 
lower slopes of Volcan de Fuego, 1140 meters, Salvin. 



360 FIELDIANA: BOTANY, VOLUME 24 

Plants arising from a tuberous root; petioles about 60 cm. long; leaf blades 
reniform in outline, glaucescent beneath, about 30 cm. wide, pedately 5-cleft, the 
segments acuminate, the 3 middle ones truncate at the base, the outermost divi- 
sions semihastate, the basal lobe rounded; peduncles shorter than the petioles; 
spathes 17-20 cm. long, the tube ventricose-tubular, the limb lanceolate, acute; 
spadix about 12 cm. long, the pistillate portion cylindric, the staminate subclavate; 
anthers 3-4. 

Xanthosoma robustum Schott, Oesterr. Bot. Wochenbl. 3: 
370. 1853. X. roseum Schott, Oesterr. Bot. Zeitschr. 8: 178. 1858. 
Quequesque; Ququeshque; Marac (Coban, Quecchi). Figure 61. 

Wet soil, usually in marshy or boggy ground, in the open, in 
thickets, or in thin forest, 900 meters or lower, most plentiful at low 
elevations; Alta Verapaz; Izabal; Jalapa; Jutiapa; Santa Rosa; Es- 
cuintla; Chimaltenango; Huehuetenango; San Marcos; Quezalte- 
nango; Retalhuleu. Honduras to Costa Rica. 

Plants very large and conspicuous, the caudex usually procumbent, often 1-4 
meters long and 10-20 cm. thick; petioles 40-180 cm. long or much longer, vaginate 
to the middle; leaf blades sagittate-ovate, often two meters long but usually shorter, 
short-cuspidate, the basal lobes half as long as the anterior one, the blades glabrous, 
grass green above, pale beneath; tube of the spathe 6-10 cm. long, 6 cm. broad, 
oblong-ovoid, greenish, the limb twice as long, ovate-lanceolate, acute, greenish 
or whitish outside, white or pink within, 12-25 cm. long and 10-25 cm. wide; spadix 
stipitate 8 mm., slightly shorter than the spathe, the pistillate portion 3-4.5 cm. 
long, almost 2 cm. thick. 

Called "quiscamote," "quiscamo," and "quiscamotillo" in Hon- 
duras, and "capote" in Chiapas. Los Quequexques is a caserio of the 
Department of Guatemala, and the vernacular name of the species 
appears in other local geographic names. The plant grows most often 
in boggy soil near stream banks, where it sometimes forms wide colo- 
nies. Its large leaves make it very conspicuous, and the large spathes 
also are showy and rather handsome. The long thick trunks, which 
lie for part of their length flat on the ground, distinguish the plant 
from all other Central American Araceae. 

This species often is planted for ornament in gardens and parks. 
The huge leaves are used as protection against sudden showers. The 
young leaves are said to be cooked and eaten in Guatemala, but the 



FIG. 61. Xanthosoma robustum. a, Habit (X Y%). b, Longitudinal section 
through pistillate flower (X 12). c, Inflorescence (X %). d, Leaf blade and 
portion of petiole (X Ve). e, Spadix with portion of spathe removed (X 1 A}- 
f, Portion of surface of pistillate spadix (X 2^)- ff, Staminate flowers, lateral 
view (X 2K). h, Portion of surface of staminate spadix (X 2^)- i, Pistillate 
flower, lateral view, showing position of ovules (X 12). 




361 



362 FIELDIANA: BOTANY, VOLUME 24 

roots are regarded as poisonous. The boiled leaves are reputed to be 
given to nursing mothers to eat in order to stimulate the natural milk 
supply. The crude milky sap is used in some areas as a substitute for 
sulfur to coagulate wild rubber (castilla) in the fabrication of rain- 
coats. 

Engler and Krause suggested that X. roseum may be only a vari- 
ety of X. robustum (Das Pflanzenreich IV. 23E. 47. 1920), and re- 
cently Matuda (Las Araceas Mexicanas, in Anal. Inst. Biol. 24: 117. 
1954) indicated his belief that X. roseum was probably synonymous 
with X. robustum. As the supposed differences in Engler and 
Krause's key (loc. cit. p. 42) between the two taxa do not appear 
to hold, as presently available material, at least, indicates, we are 
using the older name for the Guatemalan material, as suggested by 
Dr. Birdsey. 

r- 

Xanthosoma violaceum Schott, Oesterr. Bot. Wochenbl. 3: 
370. 1853. Coco (Izabal); Quequesque; Quequeshque; Badu (Puerto 
Barrios) ; Malanga (Izabal) ; Cascamote, Caxcamote (Quezaltenango) ; 
Ox (Coban, Quecchi) ; Quiscamote. 

Moist or wet thickets or forest, 1900 meters or lower; Alta Vera- 
paz; Izabal; Chiquimula; Santa Rosa; Sacatepe"quez; Retalhuleu; 
Quezaltenango; cultivated commonly in some regions, especially in 
Alta Verapaz and Izabal. Southern Mexico to Panama; West 
Indies. 

Rhizome hypogaean, short and thick; petioles 30-70 cm. long, long-vaginate; 
leaf blades sagittate-oblong-ovate, 20-50 cm. long and 15-40 cm. wide or often 
larger, paler beneath, glabrous, apiculate-acuminate, the basal lobes short, sub- 
triangular, obtuse, the sinus acute, open; peduncles 15-20 cm. long; tube of the 
spathe 10 cm. long, 3.5-4 cm. broad, oblong, glaucous, often tinged with violet or 
dark purple, the limb oblong-lanceolate, yellowish white, 15-20 cm. long, 6-7 cm. 
wide; pistillate portion of the spadix whitish, 4 cm. long, 2 cm. thick, the fertile 
staminate part 15 cm. long, the sterile portion 4 cm. long. 

Called "malanga" and "quiscamote" in Honduras; "tiquisque" 
in Costa Rica; "oto" in Panama. The fleshy roots of cultivated 
plants are cooked and eaten. The plant is much cultivated for food 
in Alta Verapaz and Izabal, and to a lesser extent in many other 
parts of the country, chiefly in the lowlands. The roots are poison- 
ous when raw, but the poisonous properties (probably due to the 
presence of irritating crystals) are destroyed by cooking. A plant 
with leaves conspicuously tinged with dark purple, probably a form 
of this species, is planted for ornament in gardens, especially in the 
Orient. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 363 

ZANTEDESCHIA Sprengel. Calla 
Reference: Engler, Pflanzenreich IV. 23Dc: 61. 1915. 

Terrestrial plants with thick rhizomes, the leaves several, long-petiolate, the 
petioles spongy, the blades mostly sagittate or hastate; peduncles long, equaling 
or exceeding the leaves; spathes large, white or yellow, rarely pink, the tube wither- 
ing and persistent, the blade open and spreading, recurved and cuspidate at the 
apex; flowers unisexual, naked; staminate flowers with 2-3 stamens, the anthers 
cuneate-quadrate, subcompressed, sessile, the thick connective truncate at the 
apex, the cells oblong, extrorse, opening by a vertical pore; pistillate flowers 1-5- 
gynous, the ovary short-ovoid, attenuate to a short style, 1-5-celled; ovules usually 
4 in each cell, biseriate, anatropous; stigma hemispheric-discoid; berries obovoid 
or subglobose, 1-5-celled, the cells 1-2-seeded; seeds ovoid, the testa longitudi- 
nally striate. 

Eight species are recognized by Engler, all native in southern 
Africa. 

Zantedeschia aethiopica (L.) Spreng. Syst. Veg. 3: 715. 1826. 
Calla aethiopica L. Sp. PI. 968. 1753. Pseudohomalomena pastoensis 
A. D. Hawkes, Madrono 11: 147. fig. 1. 1951. Cartucho; Gala. 

Grown in all regions of Guatemala for ornament, in some local- 
ities in quantity in order to sell the flowers, which are abundant in 
most of the markets; well naturalized locally in Alta Verapaz and at 
Santa Maria de Jesus in Quezaltenango, and noted as growing in 
thickets remote from dwellings in San Marcos. Native of South 
Africa. 

The common calla is too well known to need detailed description. 
In Guatemala as in most other regions the "flowers" are employed 
principally for coronas and other funeral decorations, for which they 
have long been favored. They are much used also for decorating 
altars and shrines, and are liked particularly because water does not 
have to be changed often for the cut flowers. Nowhere in Guatemala 
is the plant so well naturalized as on the humid slopes of the Volcano 
of Irazu in Costa Rica, where wide hedges of them line the lanes. 
The plants thrive best in very wet places, especially in marshes or 
swampy meadows, and they have thoroughly established themselves 
above the electric plant at Santa Maria de Jesus, where they grow 
in shallow water with native plants characteristic of such habitats. 
There is grown in Guatemala also, but rarely, the yellow calla, Zante- 
deschia Elliottiana (Knight) Engler, distinguished by having bright 
yellow rather than white spathes, and perhaps also the spotted calla, 
Z. albo-maculata (Hook, f.) Baillon, with white or cream-colored 
spathes, the leaves conspicuously spotted or streaked with white. 



364 FIELDIANA: BOTANY, VOLUME 24 

LEMNACEAE. Duckweed Family 1 

References: Hegelmaier, Systematische Ubersicht der Lemnaceen, 
Bot. Jahrb. Engler 21: 268-305. 1895; Thompson, A revision of the 
American Lemnaceae occurring north of Mexico, Rept. Mo. Bot. 
Gard. 9: 21-42. 1897; Bravo, Las Lemnaceas del Valle de Mexico, 
Ann. Inst. Biol. 1: 7-32. 1930. 

Small aquatic plants, floating free, beneath or upon the surface of the water, 
each plant consisting of a disk-shaped or elongate, fleshy or membranaceous modi- 
fied stem (frond); frond green, loosely cellular, with or without one or more roots; 
asexual vegetative reproduction by lateral branching, the branches soon separat- 
ing; inflorescence consisting of one or more naked monoecious flowers borne on the 
edge or upper surface of the frond; flower consisting of a single stamen or of a 
single flask-shaped pistil; anther 2- or 4-celled, the pollen grains spherical and 
minutely barbellate; ovary sessile, 1-celled, containing 1-6 erect or horizontal, 
anatropous to orthotropous ovules; fruit a 1-6-seeded utricle. 

The members of this family are the smallest of the flowering 
plants. Of the four genera recognized in the family, three are found 
in Guatemala. 

Fronds without roots; one reproductive pouch Wolffia 

Fronds with one or more roots; reproductive pouches two. 

Root solitary Lemna 

Roots several Spirodela 



LEMNA L. 

Fronds disk-shaped or elongate, with a central nerve and sometimes with 2 or 
4 lateral nerves, each frond with a single root proliferous from a cleft in the margin 
toward the base; flowers arising from a cleft in the margin of the frond, usually 3 
together surrounded by a spathe; 2 of the flowers staminate, consisting of 1 stamen 
only, the other flower pistillate, consisting of a simple pistil; filament slender; an- 
ther 2-celled, didymous, the cells transversely dehiscent; ovary 1-celled; style and 
truncate or funnel-shaped stigma simple; ovules and seeds 1-7. 

About 10 species, widely distributed. Besides the species listed 
here, one other is reported for Central America. 

Fronds symmetric or almost symmetric. 

Plants deep green, thickish, convex on both surfaces, oblong-obovate, obscurely 
3-veined, cavernous throughout, appearing medium-thick when pressed; 
margins thick L. minor 

Plants mostly paler green, the lower surface nearly flat, oblong-elliptic, obscurely 
1-nerved, cavernous in the middle portions only, appearing membranous 
when pressed; margins thin L. minima 

1 The Guatemalan specimens have been studied and identified by Profesora 
Maria G. de Bracco of Argentina. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 365 

Fronds asymmetric. 

Frond distinctly or indistinctly 3-nerved, obliquely obovate to oblong obovate, 
medium thick, deep green; root sheath with lateral wing appendages. 

L. paucicostata 

Frond obscurely 1-nerved or nerveless, oblong to obovate oblong, thin, pale 
green; root sheath unappendaged L. valdiviana 

Lemna minima Philippi, Linnaea 33: 239. 1864. 

Ponds and spring-fed streams, 1500-3100 meters; Totonicapan; 
Huehuetenango; Quezaltenango. United States; Mexico; Costa 
Rica; South America. 

Plants solitary or persisting in groups of 2-4; fronds symmetric or with the 
abscission scar slightly to one side of the median line, oblong to elliptic, mostly 
1.5-2.5 mm. long, 0.9-1.5 mm. broad, the apex rounded, obscurely 1-nerved or 
nerveless, with a row of papules along the midnerve, the lower surface flat or slightly 
convex, the upper surface usually conspicuously convex, cavernous in the middle 
portion only, the peripheral portion thin-margined, membranous when pressed; 
root sheath thin, cylindric, unappendaged; spathe open; pistil short-clavate, with 
concave stigma; ovule solitary; fruit elongate, erect; seed oblong, pointed, about 
16-ribbed, with many transverse striations. 

Lemna minor L. Sp. PI. 970. 1753. 

Small pools in streams, bogs, and ponds, or sometimes on moist 
slopes below waterfalls, 500-1700 meters or less; Pet&i; Zacapa; Chi- 
quimula; Jutiapa; Sacatepe*quez; Escuintla; Suchitepe'quez. United 
States; Mexico; El Salvador; South America; Old World. 

Plants solitary or 2-8 attached by the persistent internodes; fronds mostly 
symmetric, or in fruit slightly asymmetric, suborbicular to elliptic-obovate, 2-4 
mm. long, 1.5-3 mm. broad, obscurely 3- veined, rarely 4-5- veined, with a row of 
papules along the midnerve of the upper surface, both surfaces convex, cavernous 
throughout, the peripheral portion thick-margined, appearing medium-thick when 
pressed, the upper surface smooth and glistening, deep green, the lower surface 
sometimes tinged with red or purple; root sheath long, cylindric, unappendaged; 
spathe sac-like, with a small cleft opening which is irregularly pierced by the devel- 
oping floral parts; pistil clavate, with a short style and stigma; ovule solitary, 
amphitropous; fruit a symmetric subturbinate wingless utricle, projecting about 
one-third beyond the edge of the plant; seed oblong-ovate, with prominent rounded 
hilum, 12-15-ribbed, with many transverse striations. 

The Maya name of Yucatan has been reported as "Ximha." 

Lemna paucicostata Hegelm. ex Engelm. in Gray, Man. Bot. 
681. 1868. Figure 62, a-d. 

Pools, small streams, swamps, sometimes around thermal springs, 
and occasionally on moist faces of cliffs, occurring from sea level to 



366 FIELDIANA: BOTANY, VOLUME 24 

2800 meters; Pete*n; Izabal; Chiquimula; Jutiapa; Zacapa; Guate- 
mala; Sacatepe"quez; Escuintla; Retalhuleu; Quiche". United States; 
Mexico; El Salvador; Honduras; Costa Rica; South America. 

Plants solitary or occurring in groups of 2-6; fronds strongly asymmetric, 
rarely nearly symmetric in robust sterile specimens, obliquely obovate to oblong- 
obovate, 2-3.5 mm. long, 1.2-2.5 mm. broad, distinctly or indistinctly 3-nerved, 
the apical dorsal papule usually prominent and frequently a row of smaller ones 
along the midnerve, cavernous throughout, medium-thick; root sheath with lateral 
longitudinal wing appendages; spathe open; ovule solitary, obliquely orthotropous; 
fruit asymmetric, ovoid to oblong, ending in a prominent style; seed ovoid to 
obliquely oblong, slightly compressed, oblique in the utricle, the thick seed coat 
prominently 12-16-ribbed, with many transverse striations. 

This species is often considered synonymous with L. perpusilla 
Torr., but Profesora de Bracco considers the two distinct. 

A collection from the Department of Suchitepe"quez (Steyermark 
47697) has been referred by Profesora de Bracco to L. paucicostata 
var. membranacea Hegelm. 

Lemna valdiviana Philippi, Linnaea 33: 239. 1864. L. cyclo- 
stasa (Ell.) Chev. Fl. Par. 2: 256. 1827. Figure 62, e, f. 

Reported by Hegelmaier from Atitlan, Solola, Bernoulli (as L. 
valdiviana var. abbreviata) ; Quezaltenango; San Marcos. United 
States; Mexico; Panama; West Indies; South America. 

Plants solitary or in groups of 2-8; fronds usually somewhat asymmetric, ob- 
long to obovate-oblong, 2.3-4.5 mm. long, 0.7-1.5 mm. broad, obscurely 1-nerved 
or nerveless, thin, without papules, cavernous in the middle portions only; root 
sheath large, thin, cylindric, unappendaged; rootcap strongly curved; spathe open, 
reniform; style relatively long, straight or sometimes curved; ovule solitary, 
obliquely orthotropous; fruit slightly asymmetric, ovoid-oblong, about half the 
length of the frond; seed oblong-ovoid, the thick seed coat 12-19-ribbed, with 
many transverse striations. 

SPIRODELA Schleiden 

Fronds orbicular to oblong or obovate, 5-15-nerved, each frond with 2-16 
fascicled roots containing a single bundle of vascular tissue, the stipe attached 
peltately to the frond back of and below the basal margin; flowers arising from 2 
triangular vegetative reproductive pouches which open as clefts in either margin of 



FIG. 62. a-d, Lemna paucicostata: al, Sterile plant (X 14); a2, Plant with 
fruit (X 14); a3, 4, Seeds (X 7). b, Broad fronds (X 14; Standley 89331). c, 
Narrow frond (X 14; Steyermark 46269). d, L. p. var. membranacea (X 14; 
Steyermark 47697). e, f, L. valdiviana (X 14; e, Steyermark 36351; f, Steyermark 
3388^). g, Wolffia Welwitschii: gl, Sterile plant (X 14); 02, Plant with flowers 
(X 14). 






a 












367 



368 FIELDIANA: BOTANY, VOLUME 24 

the basal portion of the frond, usually 3 together surrounded by a sac-like spathe; 
2 of the flowers staminate with filaments curving upward from the margin of the 
frond; anthers 2-celled, longitudinally dehiscent; one flower pistillate; ovary with 
2 anatropous ovules; fruit rounded-lenticular, the margins winged. 

About 3 species are recognized, one occurring in Asia and Austra- 
lia and another in South America. Only the following is known from 
Central America. 

Spirodela polyrhiza (L.) Schleid. Linnaea 13: 392. 1839. Lemna 
polyrhiza L. Sp. PL 970. 1753. 

Margins of lakes and swamps along streams and at the base of 
wooded bluffs, 400-500 meters; Pete"n; Chiquimula; Jutiapa; Guate- 
mala; Retalhuleu. Widely distributed in both hemispheres. 

Plants solitary or in colonies of 2-5 attached by the short persistent internodes; 
fronds slightly asymmetric, suborbicular to suborbicular-obovate, 2.5-9.5 mm. 
long, 2.2-6.5 mm. broad, 5-15-nerved, each nerve with one vascular bundle, the 
upper surface bluish green or pale yellow-green above, the lower surface deep pur- 
plish red; roots 3-16 on each plant, one vascular bundle in each root; rootcap large 
and thin; spathe sac-like, opening at the upper end; pistil flask-shaped; ovules 2, 
anatropous, or frequently one and then amphitropous; fruit a rounded utricle 
with slightly winged margins; seed slightly compressed, smooth. 

WOLFFIA Horkel 

Fronds globose or oblong-ovoid, fleshy, rootless, nerveless, proliferating from 
a cleft or opening at the basal end; flowers arising from the upper surface of the 
frond, breaking through the loosely cellular tissue, the spadix consisting of 2 flowers 
without a spathe, one flower staminate, the other pistillate; staminate flower of 
one stamen with a 1-celled anther opening by a slit across the top, the two halves 
opening as valves; pistillate flower of a globular ovary with a short style, de- 
pressed stigma, and single orthotropous ovule; fruit a spherical utricle. 

This genus consists of the simplest and smallest flowering plants 
known, the individual plants appearing as tiny specks on the surface 
of the water. About 8 species, mostly in tropical or subtropical re- 
gions; only the following are known from Central America. 

Fronds 1-flowered, 0.5-1 mm. long, 0.5-0.8 mm. broad, upper surface convex. 

W. columbiana 
Fronds 2-flowered, 0.5-0.6 mm. long, 0.3-0.4 mm. broad, upper surface flat. 

W. Welwitechii 

Wolffia columbiana Karst. Bot. Unters. 1: 103. 1865. 

Guatemala (Moran, 1205 meters, Kellerman 6630) ; also reported 
by Hegelmaier from Atitlan, Solola (Bernoulli). Mexico to South 
America. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 369 

Frond symmetric, elliptic to globose, 0.5-1 mm. long, 0.5-0.8 mm. broad, the 
upper surface convex, slightly exposed to the air, with usually about 3 small incon- 
spicuous papules in a row along the median line, obtuse at each end, not punctate; 
single-flowered. 

The Maya name of Yucatan for this plant is "iximha." Although 
we have not seen the specimen, it is probable that the Wolffia punc- 
tata Griseb. reported by Sereno Watson from Rio Chocon, Izabal, 
should be placed with W. columbiana. 

Wolffia Welwitschii Hegelm. Seem. Journ. 114. 1865. Fig- 
ure 62, g. 

Jutiapa (Lago Guija, southeast of Asuncion Mita, 470-500 me- 
ters, Steyermark 31812). Cuba; Venezuela; tropical Africa. 

Frond symmetric, flat, thin, elliptic, with many pigment cells in the epidermis, 
0.5-0.6 mm. long, 0.3-0.4 mm. broad; two flowers present on a frond; anther 0.1 
mm. long; pistil 0.25 mm. high. 

MAYACACEAE 

Small mosslike herbs of wet soil, sometimes floating in water, prostrate or rep- 
tant, branched, the stems densely leafy; leaves short, linear-filiform, spreading in 
all directions; flowers perfect and regular, the pedicels 1-flowered, solitary in the 
leaf axils, bracteate at the base and often bearing 2 small bractlets; perianth infe- 
rior, biseriate, each whorl 3-parted, the outer calyx-like, the 3 sepals lanceolate, 
distinct; petals 3, distinct, obovate or rounded, subequal, spreading, in bud imbri- 
cate; stamens 3, hypogynous, the filaments short and filiform; anthers erect, basi- 
fixed, oblong, the 2 cells parallel, partitioned within, short-dehiscent at the apex; 
ovary sessile, superior, 1-celled, with 3 parietal placentae; style filiform, simple, 
the small stigma terminal, entire; ovules numerous on each placenta, biseriate, 
orthotropous; capsule surrounded by the persistent calyx and stamens, ovoid, 
membranaceous, dehiscent by 3 more or less distinct valves; seeds ovoid or glo- 
bose, with basal hilum, terminated by a small tubercle, the testa longitudinally 
striate and usually tuberculate-rugulose; endosperm farinaceous. 

The family consists of a single genus with about 7 species in the 
warmer regions of America. Only one species is known from Cen- 
tral America. 

MAYACA Aubl. 
Mayaca fluviatilis Aubl. PI. Guian. 42. 1775. Figure 63. 

In bogs or sometimes in flowing water or shallow ponds, at 1500 
meters or less; British Honduras and probably in Pete"n; Alta Vera- 
paz; Jalapa. Southeastern United States; Honduras; Costa Rica; 
Panama; West Indies; South America. 



370 



FIELDIANA: BOTANY, VOLUME 24 




FIG. 63. Mayaca fluviatilis. a, Habit (X ^). b, Portion of leafy stem with 
flower bud (XI). c, Fruit with sepals (X 7). d, Expanded flower (X 7). e, 
Longitudinal section through ovary (X 18). 



A small plant, the stems often only 2-3 cm. long but usually longer and some- 
times as much as 40 cm., very densely leafy; leaves lance-linear to filiform, 4-12 
mm. long, soft, usually very densely crowded at the tips of the branches, spreading, 
bright green; pedicels equaling or usually shorter than the leaves, often recurved in 
age; sepals green, 3-4 mm. long; capsule 4-5 mm. long. 

This has been reported from Central America as M. Aubletii 
Michx., which is apparently a different species. In general appear- 
ance it much resembles some of the species of Lycopodium, so much 
so that at first glance it is difficult to decide which genus is repre- 
sented. The white flowers are very small and inconspicuous. The 
plant is to be expected in the savannas of Izabal. 



XYRIDACEAE 

Perennial or rarely annual herbs, often cespitose; leaves radical, rosulate or 
crowded, sometimes distichous, generally linear; scapes erect, rushlike, simple, 
naked; flowers crowded in a terminal, globose or ovoid head, solitary and sub- 
sessile within imbricate, coriaceous or rigid bracts; flowers perfect, slightly irreg- 
ular; perianth inferior, corolloid, simple or double; sepals 3, the middle one large, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 371 

petaloid, surrounding the corolla, caducous or sometimes absent; corolla tube slen- 
der, short or elongate, the 3 lobes equal, spreading, broad; perfect stamens 3, in- 
serted near the base of the lobes, shorter than the lobes; filaments short, filiform; 
anthers oblong, dorsifixed, the 2 cells parallel, or divergent at the base, dehiscent 
by a longitudinal slit; staminodia 3 and alternate with the corolla lobes, or none; 
ovary sessile, 1-celled or imperfectly 3-celled, the 3 placentae erect from the base 
of the ovary or usually more or less adnate to the walls; style terminal, simple, or 
with 3 branches stigmatiferous at the apex; ovules numerous in each cell or some- 
times few, 2-seriate, orthotropous; capsule included in the marcescent corolla tube, 
3-valvate; seeds numerous, ovoid, bearing a small tubercle at the apex, the testa 
vertically striate; endosperm farinaceous. 

Only two genera are known, the other, Abolboda, with very few 
species, these all in tropical America, but not reaching Central 
America. 

XYRIS L. 

Reference: A. G. 0. Malme, Die amerikanischen Spezies der Gat- 
tung Xyris L., Untergattung Euxyris (Endlicher), Arkiv Bot. 13, 
no. 8: 1-32. 1913; N. Amer. Fl. 19: 3-15. 1937. 

Annual or perennial, tufted herbs; leaves linear, equitant; scapes simple, ter- 
minated by a solitary dense head, usually with a single basal sheath; flowers yellow, 
almost regular, in globose to ovoid heads; blades of the petals cuneate-obovate, 
spreading; stamens alternating with staminodia, the filaments short, flattened; 
staminodia usually bifid at the apex, the branches terminated by tufts of small 
hairs; ovules few or numerous. 

About 100 species, widely distributed in tropical regions. One 
other is known from Central America. 

Flower heads small, 5-7 mm. long; leaves 4-6 cm. long, 1-1.5 mm. wide; keel of 
the lateral sepals ciliate-scabrous or dentate, especially at the middle, rarely 
toward the apex X. subnavicularis 

Flower heads large, 8-35 mm. long; leaves 15-80 cm. long, 3-10 mm. wide; keel 
of the lateral sepals lacerate-dentate, fimbriate, or villous above. 

Heads narrowed and subacute at the apex; keel of the lateral sepals ciliate-sca- 
brous; scapes multistriate or multicostate below X. ambigua 

Heads broad and rounded at the apex; keel of the lateral sepals dentate, lacerate, 
or lacerate-fimbriate; scapes 1-2 costate X. Jupicai 

Xyris ambigua Beyrich ex Kunth, Enum. PI. 4: 13. 1843. 

Savannas or pine ridges, sometimes about ponds, at or little 
above sea level; British Honduras; southeastern United States; Cuba. 

Plants tall, the leaves 15-40 cm. long, 3-6 mm. wide, subobtuse, the margins 
scabrous or rarely smooth; sheath of the leaf about as long as the blade, slightly 
dilated and opaque below, pale rusty at the base; scapes 50-90 cm. tall, 1.5-2 mm. 
thick, subterete, bicostate above, multicostate below, scabrous on the angles; pe- 
duncular sheath 10-20 cm. long; heads many-flowered, ellipsoid or ovoid, 10-25 



372 FIELDIANA: BOTANY, VOLUME 24 

mm. long, 5-7 mm. thick, the outer sterile bracts 2-2.5 mm. long, ovate-deltoid, 
acute or subobtuse, the fertile bracts obovate-elliptic, 7-8.5 mm. long, 5-6 mm. 
wide, rounded at the apex, entire or somewhat lacerate at the apex, stramineous, 
fulvous, or pale-ferruginous, lustrous, with a deltoid or ovate dorsal area about 
2 mm. long; lateral sepals lanceolate, 6 mm. long, 1.5 mm. wide, acute; keel broad, 
scabrous-ciliate from about the middle to the apex; seeds ellipsoid, 0.4-0.5 mm. 
long. 

Xyris Jupicai L. Rich. Act. Soc. Hist. Nat. Paris 1: 106. 1792. 
X. communis Kunth, Enum. PI. 4: 12. 1843. Figure 64. 

In marshes or bogs, sometimes in sandy soil along streams, 1700 
meters or lower; Alta Verapaz; Izabal; Chiquimula; Jalapa; Jutiapa; 
Santa Rosa. Southern Mexico; British Honduras; Honduras; south- 
ern United States; West Indies; South America. 

Leaves 12-40 cm. long, 3-5 mm. wide, acute, smooth; sheath equaling the leaf 
blade or only half as long, fulvous or somewhat ferruginous below, somewhat di- 
lated at the base; scapes usually 30-65 cm. tall, 1-2 mm. thick, 1-2-costate, smooth, 
the sheath usually 8-15 cm. long, ferruginous or chestnut and shining beneath; 
heads many-flowered, ovoid or ellipsoid, 8-13 mm. long, 5-8 mm. thick, the outer 
bracts ovate-elliptic, 2-3 mm. long, rounded at the apex, the floriferous bracts obo- 
vate, 5-7 mm. long, 3.5-4.5 mm. wide, entire, fulvous or ferruginous, somewhat 
lustrous, with a grayish green or green, ovate or elliptic dorsal area 2 mm. long; 
lateral sepals narrowly spatulate-linear or lanceolate, 3.5-5 mm. long, 0.6 mm. 
wide, acute; keel narrow and entire below, broader above and lacerate-dentate or 
shortly lacerate-fimbriate from the middle to the apex; seeds ellipsoid, 0.5 mm. 
long. 

Xyris subnavicularis Malme, Arkiv Bot. 13, no. 8: 15. 1913. 

Wet soil, at or little above sea level; British Honduras; type from 
Stann Creek, John Robertson; also at All Pines. 

Leaves linear, 4-6 cm. long, 1-1.5 mm. wide, subobtuse, scabrous-ciliate on 
the margins, smooth elsewhere; leaf sheath about equaling the blade, pale-ferrugi- 
nous, lustrous at the base; scapes 10-15 cm. tall, 0.5-0.7 mm. thick, terete or nearly 
so, bicostate, especially above, scabrous on the edges, elsewhere smooth, the sheath 
about 4 cm. long; head few-flowered, ellipsoid, obovoid, or subglobose, 5-7 mm. 
long, the outer sterile bracts ovate, 3 mm. long, acute or subacute, carinate, the 
floriferous bracts elliptic, 4-5 mm. long, 3 mm. wide, rounded at the apex, entire, 
carinate above, ferruginous or fulvous, without a distinct dorsal area; lateral sepals 
lanceolate, falcate, about 3.5 mm. long and 0.8 mm. wide, acute; keel narrow, sca- 
brous-ciliate in the middle and often also toward the apex. 



FIG. 64. Xyris Jupicai. a, Corolla (X 2%). b, Habit showing leaves and 
lower part of scape (X 14)- c, Bract (X 6). d, Habit of inflorescence and upper 
part of scape (X %). e, Lateral sepal (X 10). 




dl 




373 



374 FIELDIANA: BOTANY, VOLUME 24 

ERIOCAULACEAE. Pipewort Family 

References: Harold N. Moldenke, Eriocaulaceae, N. Amer. Fl. 19: 
17-50. 1937; The Eriocaulaceae of the Yucatan Peninsula, Carnegie 
Inst. Wash. Publ. 522: 139-147. 1940. 

Mostly perennial herbs of wet soil, sometimes growing in water, the stems usu- 
ally short; leaves mostly basal and tufted, narrow and grass-like, generally crowded; 
flowers monoecious, rarely dioecious, capitate, in terminal, solitary or umbellately 
aggregate, involucrate heads, borne on usually long and slender, scapose peduncles, 
these vaginate at the base; flowers regular, numerous, very small, sessile or short- 
pedicellate on a receptacle, each borne in the axis of a scarious scalelike bractlet; 
staminate and pistillate flowers mixed together, the staminate usually in the center; 
perianth scarious or membranous, 2-3-parted; stamens as many as the outer peri- 
anth segments and alternate with them or twice as many; filaments distinct, the 
anthers small, 2-4-celled, opening by longitudinal slits, introrse; ovary superior, 
2-3-celled, the style terminal; stigmas 2-3, simple or lobate; ovules solitary and 
pendulous in each cell, orthotropous; capsule 2-3-celled, membranous, loculicidally 
dehiscent; endosperm copious. 

Nine genera with many species, widely dispersed in tropical re- 
gions of both hemispheres, a few species in temperate regions. The 
family is best represented in Brazil, especially in the region of Minas 
Geraes. Only the following genera are represented in Central Amer- 
ica. The plants constitute a difficult group, with microscopic flowers 
that can be dissected with difficulty. The following treatment is 
copied from those by Moldenke. The family is surprisingly well rep- 
resented in the pine forests of British Honduras, but the Guatemalan 
species are very few indeed. 

Plants with elongate leafy stems, the stems floating on or submerged in water; 

peduncles axillary Tonina 

Plants acaulescent or essentially so, usually terrestrial, the leaves all at the base of 
the scape, the peduncles terminal, or at least appearing so. 

Stamens 4 or 6, twice as many as the outer perianth segments Eriocaulon 

Stamens 2-3, as many as the outer perianth segments. 

Inner perianth segments of the pistillate flowers free Paepalanthus 

Inner perianth segments of the pistillate flowers connate at the middle. 

Syngonanthus 

ERIOCAULON L. Pipewort 

Plants mostly perennial, the stems very short in Central American species; 
leaves often numerous at the base of the scape, linear or nearly so, amplexicaul; 
flowers 2-3-parted, the staminate mixed with the pistillate or segregated in sepa- 
rate heads; perianth almost always double; sepals of the staminate flower free at 
the base or more or less connate; petals 2-3, united below into a tube, the lobes 
usually bearing a small black gland on the inner surface near the apex; stamens 
twice as many as the sepals, exserted, the anthers 4-celled, mostly black; pistillate 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 375 

flowers with free or rarely connate sepals, the petals free or rarely none, usually 
black-glandular inside below the apex; stigmas 2-3, simple. 

About 360 species, mostly in tropical regions, and most numerous 
in Brazil. Three other species are known from southern Central 
America. 

Pistillate flowers without glands E. bilobatum 

Pistillate flowers with glands. 

Staminate and pistillate flowers 2-parted ; stamens 4; stigmas 2; ovary 2-ovulate. 
Mature flower heads white-villous at the apex, 4.5-7 mm. in diameter. 

E. Kinlochii 

Mature flower heads blackish, glabrous, 1.5-3 mm. in diameter. 

E. fuliginosum 

Staminate and pistillate flowers 3-parted ; stamens 6; stigmas 3; ovary 3-ovulate. 

Stems elongate, floating E. Schippii 

Stems short, not floating. 

Petals of the pistillate flowers black-glandular E. Ehrenbergianum 

Petals of the pistillate flowers not glandular E. Williamsti 

Eriocaulon bilobatum Morong, Bull. Torrey Club 19: 226. 1892. 

In wet mud bordering a swamp, about 1000 meters; Jutiapa 
(plains between Agua Blanca and Amatillo, Steyermark 30405). 
Jalisco, Mexico. 

Plants probably annual, acaulescent or nearly so; leaves rather numerous, nar- 
rowly linear, 1.5-5.5 cm. long, about 1.2 mm. wide, subulate at the apex, 2-3- 
nerved, glabrous; peduncles 2-8 and 1.5-11 cm. long, 5-costate, glabrous; flower 
heads blackish, ovoid-globose, 2-3.5 mm. in diameter; involucral bracts grayish, 
oblong or lance-oblong, acute or subacute, glabrous; bractlets lance-oblong, longer 
than the flowers and hiding them; Staminate sepals 2-3, connate into a glabrous, 
grayish white or stramineous spathe, this shortly 3-fid at the apex; petal tube bear- 
ing at the apex very small, slightly unequal lobes, these marked with fuscous 
glands; stamens 6; pistillate sepals 2-3, very narrowly linear, unequal, glabrous; 
petals none; stigmas 3, filiform; ovary 3-ovulate. 

Eriocaulon Ehrenbergianum Klotzsch ex Koern. in Mart. Fl. 
Bras. 3, pt. 1:491. 1863. 

Open bogs, 1500-1700 meters; Sacatepe"quez; Quiche". Mexico. 

Plants acaulescent; leaves linear or ensiform, 1-8 cm. long, 2 mm. wide, ob- 
tusely subulate at the apex, 5-10-nerved, glabrous; peduncles 1-5, much exceeding 
the leaves, 7-30 cm. long, 6-sulcate, glabrous; sheaths rather loose, about equaling 
the leaves; flower heads subglobose, 5-8 mm. in diameter, white-villous; involucral 
bracts membranous, golden brown or yellowish green, ovate, very shortly cuspi- 
date; staminate sepals 3, spathaceous-connate, olivaceous-nigrescent, spatulate- 
oblong, subacute, pilose at the apex; corolla lobes 3, short, pilose, black-glandular; 
anthers 6, black; pistillate sepals 3, olivaceous-nigrescent, broadly lanceolate, acute, 



376 FIELDIANA: BOTANY, VOLUME 24 

pilosulous dorsally, carinate; petals 3, whitish, obtuse, glandular, the upper half 
pilose on both surfaces. 

Eriocaulon fuliginosum C. Wright ex Griseb. Cat. PL Cub. 
226. 1866. 

In pine ridges, at or little above sea level; British Honduras; 
Cuba. 

Plants acaulescent; leaves spreading or recurved, narrowly or rather broadly 
linear, often ensiform, narrowed from the base to the apex, 5-9-nerved, glabrous; 
peduncles numerous, 2.5-18 cm. long, 5-7-costate, glabrous; sheaths shorter than 
the leaves; heads very small, hemispheric, blackish, 1.5-3 mm. in diameter, gla- 
brous; involucral bracts membranous, yellowish hyaline or grayish brown, broadly 
ovate or ovate-elliptic, subacute, glabrous; receptacle glabrous; staminate sepals 2, 
cuneiform-spatulate or obovate, glabrous, slightly connate at the base; petal tube 
white, the 2 lobes minute, subacute, not glandular; anthers 4, black; pistillate 
sepals 2, dark olivaceous or fuscous, obovate, acute, carinate, glabrous; petals 2, 
black or almost hyaline, oblong-spatulate or obovate, not glandular, glabrous; style 
short, the stigmas 2, longer than the style. 

This has been reported from British Honduras as E. Schiedeanum 
Koern., a species of central Mexico. 

Eriocaulon Kinlochii Moldenke, N. Amer. Fl. 19: 23. 1937. 

Known only from the type, Stann Creek Valley, Stann Creek 
District, British Honduras, J. B. Kinloch 213. 

Plants acaulescent; leaves numerous, tufted, spreading or recurved, broadly 
linear, 3-8 cm. long, 1 mm. wide, long-attenuate and filiform at the apex, 6-7- 
nerved, glabrous; peduncles mostly 1 on each plant, 5.5-14 cm. long, 3-costate, 
glabrous; sheaths loose, 1-3 cm. long; flower heads hemispheric or globose, 4.5-7 
mm. in diameter, white- villous; involucral bracts pale stramineous, elliptic-obo- 
vate, acute or obtuse, glabrous; receptacle glabrous; staminate flowers short-pedi- 
cellate, the 2 sepals hyaline, obovate, cucullate, pilose dorsally at the apex; petal 
tube whitish, the 2 lobes brown-glandular at the apex, short-pilose on both sur- 
faces; anthers 4, black; pistillate flowers on very short pedicels, the 2 sepals hya- 
line, obovate, acute, carinate, short-pilose toward the apex; petals 2, obovate, 
subacute, short-pilose toward the apex, glandular at the apex; ovary long-stipitate, 
2-celled. 

This has been reported from British Honduras as E. Benthamii 
Kunth, a species of Mexico. 

Eriocaulon Schippii Standl. ex Moldenke, N. Amer. Fl. 19: 
34. 1937. 

Known only from the type, All Pines, Stann Creek District, Brit- 
ish Honduras, at sea level, growing in shallow pools in brackish 
swamps, W. A. Schipp 647. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 377 

Stems elongate, floating or submerged, densely leafy below; leaves narrowly 
linear, capillaceous toward the apex, flat, lax, 4-8 cm. long, 0.4 mm. wide, glabrous; 
peduncles numerous, umbellately congested in groups of 6-16 at the ends of the 
stems, faintly 5-7-striate, glabrate; flower heads blackish, hemispheric, 2-3.5 mm. 
in diameter; involucral bracts blackish, ovate, obtuse, glabrous; receptacle gla- 
brous; staminate sepals 3, grayish, subacute, glabrous; petal tube white, the lobes 
very small, hyaline, black-glandular, glabrous; anthers 6, black; pistillate sepals 3, 
blackish, connate at the base, obovate, obtuse, not carinate, sparsely pilose dor- 
sally; petals 3, narrowly spatulate, black-glandular, glabrous; ovary 3-celled. 

Eriocaulon Williamsii Moldenke, N. Amer. PI. 19: 36. 1937. 
Orange Walk District, British Honduras, in clay soil in dried 
ponds, W. C. Meyer 134- 

Plants dwarf, acaulescent; leaves tufted, erect or spreading, linear, 1-4.5 cm. 
long, 1-2 mm. wide, subulate at the apex, many-nerved, glabrous; peduncles 3-4, 
rather obscurely 3-costate, 1.5-6.5 cm. long, glabrate; flower heads hemispheric or 
ovoid-conic, dark gray, 2.5-4 mm. in diameter; involucral bracts rather few, light 
stramineous, ovate, obtuse, glabrous; receptacle glabrous; staminate flowers long- 
pedicellate, the 3 sepals blackish, glabrous; petal tube pale stramineous, the lobes 
minute, glabrous; anthers 6, white; pistillate flowers pedicellate, the 3 sepals nar- 
rowly linear, glabrous; petals hyaline, narrowly linear, glabrous; stigmas 3. 



PAEPALANTHUS Martius 

Plants various in habit, the local species with short stems or acaulescent; leaves 
mostly narrow and grass-like; flower heads villous, the flowers mostly polygamous, 
2-3-parted; perianth double, involute; staminate sepals more or less connate near 
the base, the petals connate into a hollow, usually glabrous, eglandular, slightly 
2-3-lobate tube; stamens 2-3, opposite the petals, exserted, the anthers 4-celled; 
pistillate sepals usually connate at the very base; petals free, eglandular; ovary 
2-3-celled, the style usually appendaged; stigma simple or more often 2-fid. 

About 450 species, in tropical America, most numerous in Brazil. 
One other Central American species occurs in Costa Rica. 

Flower heads gray, cylindric-elongate P. Gentlei 

Flower heads dark brown, globose P. Lamarckii 

Paepalanthus Gentlei Moldenke, N. Amer. Fl. 19: 40. 1937. 

Endemic in British Honduras, at or little above sea level, growing 
in wet soil; type from Maskall, P. H. Gentle 992. 

Stems simple, 3-4 cm. long; leaves linear, 2-4.5 cm. long, 3-4 mm. wide, atten- 
uate to the acute apex, dilated and clasping at the base, sparsely strigillose or cilio- 
late; peduncles clustered at the ends of the stems, 3-8.5 cm. long, 3-costulate, 
glabrate; sheaths 10-15 mm. long; flower heads gray, cylindric-elongate, 3-7 mm. 
long, 3 mm. broad; outer involucral bracts stramineous, elliptic-obovate, acumi- 
nate, eiliate; receptacle pilose; staminate sepals 3, light brown, acute, ciliate; an- 



378 FIELDIANA: BOTANY, VOLUME 24 

thers 3, white; pistillate sepals 3, obtuse, glabrous, the petals narrowly oblong, 
ciliolate above; seeds light yellow, cancellate, the striae minutely puberulent. 

Paepalanthus Lamarckii Kunth, Enum. PL 3: 506. 1841. 

In wet soil, at or little above sea level ; British Honduras; Panama ; 
West Indies; South America. 

Stems simple, 2-8 cm. long; leaves linear-lanceolate or broadly linear, 1.5-3 
cm. long, 0.5-2.3 mm. wide, dilated and clasping at the base, narrowed to a rather 
acute or obtuse apex, multistriate, sparsely puberulent or pilose; peduncles clus- 
tered at the end of the stem, usually numerous, 1.5-7 cm. long, pilose; sheaths 
9-13 mm. long, long-pilose; flower heads dark brown, globose, 2-3 mm. in diam- 
eter, villous; involucral bracts gray-brown, obovate, subacute, densely pilose at the 
apex; receptacle pilose; staminate sepals 3, dark brown at the apex, spatulate, ob- 
tuse, ciliate at the apex; stamens 3; pistillate sepals pilose on the margins and apex; 
ovary 3-celled; seeds slightly curved, cancellate. 



SYNGONANTHUS Ruhland 

Stems usually very short; leaves mostly grass-like; flower heads pilose or sub- 
glabrate; bractlets usually none on the receptacle; flowers 3-parted, the sepals free 
or nearly so; staminate petals connate into a 3-lobate glabrous tube; anthers 4- 
celled; pistillate petals connate at or above the middle, the base and apex free. 

About 160 species, mostly in tropical America, a few in tropical 
Africa. Only the following occur in Central America. 

Peduncles glabrous or nearly so S. Oneillii 

Peduncles conspicuously pubescent. 

Involucral bracts hyaline, not colored S. Bartlettii 

Involucral bracts more or less olivaceous. 

Sheaths shorter than the leaves, densely short-pubescent; pubescence of the 
peduncles mostly appressed; bracts as much as 2.5 mm. long, glabrous. 

S. hondurensis 

Sheaths longer than the leaves, loosely long-pilose; pubescence of the peduncles 
spreading; bractlets as much as 8 mm. long, villous S. Lundellianus 

Syngonanthus Bartlettii Moldenke, Phytologia 1: 335. 1938. 

British Honduras, endemic, growing in wet flat uplands and in 
pine ridges; type from Mountain Pine Ridge, El Cayo District, 
H. H. Bartlett 11670. 

Plants 2-7 cm. high; leaves cespitose, linear, recurved and appressed to the 
ground, 3-12 mm. long, obtuse, glabrate or obscurely puberulent; sheaths hirsute; 
peduncles solitary or few, conspicuously pilose with spreading hairs; heads 2-5 mm. 
high, the involucral bracts hyaline, colorless, elliptic-lanceolate, acute or acumi- 
nate, glabrous; staminate flowers pedicellate, densely long-tomentose at the base; 
sepals 3, elliptic-obovate, 1 mm. long, acute; petals hyaline, forming a very narrow 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 379 

tube; pistillate flowers pedicellate, the sepals free, lance-ovate, 1.5 mm. long, acu- 
minate, not ciliate; petals linear-oblanceolate. 

Syngonanthus hondurensis Moldenke, Phytologia 1 : 344. 1939. 

Known only from the type, wet pine forest, three miles west of 
Boomtown, Belize District, British Honduras, H. O'Neill 8543. 

Plants 6-7 cm. high; leaves cespitose, linear, spreading, 10-16 mm. long, 1 mm. 
wide, glabrate; peduncles 4-5, very slender, white-pilose with usually appressed 
hairs; flower heads 2-6 mm. wide; involucral bracts elliptic, membranous, oliva- 
ceous with scarious margins, 1.5-2.5 mm. long, acute or acuminate, glabrous, not 
ciliate; staminate flowers pedicellate, the 3 sepals free, 1 mm. long, acute, glabrous; 
petals united to form a tube about as long as the sepals, glabrous; pistillate flowers 
pedicellate, the pedicels densely long-villous at the base with white hairs; sepals 3, 
lanceolate, 1.8 mm. long, acute, long-ciliate, glabrous; petals narrowly oblong, 
slightly shorter than the sepals, densely long-villous outside with white hairs. 

Syngonanthus Lundellianus Moldenke, Phytologia 1: 345. 
1939. 

Known only from the type, oak ridges, Broken Ridge, 8 miles 
northeast of Boomtown, Belize District, British Honduras, H. O'Neill 
8546. 

Plants 6.5-8.5 cm. high; leaves cespitose, linear, 1-1.5 cm. long, spreading, ap- 
pressed-puberulent; sheaths longer than the leaves, pilose with spreading hairs, 
acute or subacuminate; peduncles numerous, conspicuously pilose with spreading 
hairs; flower heads 2-15 mm. wide, the involucral bracts linear-oblong, often elon- 
gate to 8 mm., the outer ones or all much shorter, acute, villous on the outer sur- 
face; staminate flowers pedicellate, the pedicels villous-tomentose at the base; 
sepals 3, free, hyaline, elliptic, 1 mm. long, acuminate, glabrous, not ciliate; petals 
united into a narrow hyaline glabrous tube almost as long as the sepals; pistillate 
flowers pedicellate, the sepals elliptic, 1.5 mm. long, acute or acuminate, gla- 
brous, not ciliate; petals narrowly oblong, shorter than the sepals, appressed- vil- 
lous dorsally. 

It is stated that the seeds sometimes germinate in the heads, pro- 
ducing small plants that flower while still in the heads an unusual 
instance of vivipary. 

Syngonanthus Oneillii Moldenke, Phytologia 1: 346. 1939. 

British Honduras; known only from pinelands near Boomtown, 
Belize District, the type being H. O'Neill 8548. 

Plants 6-10 cm. high; leaves cespitose, linear, spreading and more or less re- 
curved, 1-2.5 cm. long, 0.5-1 mm. wide, abruptly acute, glabrate or laxly pilose; 
sheaths equaling or shorter than the leaves, sparsely and laxly pilose or glabrate; 
peduncles numerous, very slender, glabrous or nearly so; flower heads 3-5 mm. 
broad, the involucral bracts oblanceolate-elliptic, rather light olivaceous or brown- 
ish, abruptly acute or obtuse, glabrous; staminate flowers pedicellate, the pedicels 



380 FIELDIANA: BOTANY, VOLUME 24 

glabrous except for a few long white hairs at the base; sepals 3, about 1 mm. long, 
oblanceolate, glabrous, short-acuminate or sinuate-tridentate at the apex; petals 
united at the middle, free above and below, slightly shorter than the sepals, in- 
curved and pilose at the apex on the outer surface; pistillate flowers pedicellate, 
the pedicels long-hirsute at the base; sepals 1.2 mm. long, lance-ovate, acute, long- 
ciliate toward the apex; petals narrowly oblong, connate by their margins. 



TONINA Aublet 

Plants probably perennial, with slender elongate stems, these equally leafy 
throughout; peduncles arising in or near the leaf axils along the stem, the flowers 
very small, capitate, 3-parted, the receptacle pilose; staminate sepals connate to 
the middle, the petals connate into a short membranous, shortly 3-lobate tube; 
stamens 3, the anthers 2-celled; pistillate sepals connate at the base, the petals 
very small, free, long-pilose; stigmas 3, bifid. 

The genus consists of a single species. 

Tonina fluviatilis Aubl. PL Guian. 857. pi. 330. 1775. 

Floating in swamp, at sea level, All Pines, Stann Creek District, 
British Honduras, W. A. Schipp 693. Vera Cruz; Costa Rica; 
West Indies; South America. 

Stems often branched, sometimes a meter long, densely leafy; leaves spreading, 
lanceolate or oblong, 8-15 mm. long, 1-2.5 mm. wide, sessile and amplexicaul, 
acute, long-ciliate; peduncles numerous, 2-13 mm. long, glabrous; heads echinate- 
globose, 4-8 mm. in diameter; involucral bracts ovate or obovate, cuspidate, gla- 
brous except at the base and apex; staminate sepals broadly obovate, abruptly 
acute; petals of the pistillate flower linear. 



BROMELIACEAE. Pineapple Family 
By LYMAN B. SMITH 

References: Mez, Bromeliaceae, Pflanzenreich, fam. 32: 1-667. 
1934-5. L. B. Smith, Bromeliaceae, N. Amer. Flora 19: 61-228. 
1938. L. B. Smith & C. L. Lundell, Bromeliaceae Yucatan Penin- 
sula, Bot. Maya Area 16: 105-136. 1940. 

Herbs in all the Guatemalan species, mostly epiphytic or saxicolous; leaves 
spirally arranged, usually basal, simple, entire or spinose-serrate, at least in youth 
bearing peltate scales serving to collect and hold moisture; inflorescence simple or 
compound, usually bearing brightly colored bracts; flowers perfect or functionally 
dioecious; perianth heterochlamydeous, the segments free or variously joined, 
trimerous; stamens 6, filaments free or joined to the petals or to each other; ovary 
superior to inferior, 3-celled; fruit capsular or baccate; seeds naked, winged, or 
plumose; embryo small, at the base of the mealy endosperm. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 381 

About 45 genera and 1600 species, indigenous to tropical and sub- 
tropical America except for a single African species. 

Ovary wholly or partly superior; fruit capsular; seeds appendaged. 
Seeds with entire appendages; leaves often spinose-serrate; ovary often only in 

part superior; plants usually terrestrial. 
Sepals not over 5 mm. long; flowers small and numerous. 

Flowers perfect; leaves thin, entire; plants slender, low Lindmania 

Flowers functionally dioecious; leaves thick, spinose-serrate; plants coarse, 

1-2 meters high Hechtia 

Sepals 15-50 mm. long; flowers conspicuous Piicairnia 

Seeds plumose; leaves always entire; ovary nearly or quite superior; plants 

chiefly epiphytic. 

Appendage of the seed basal, straight at maturity. 
Petals nearly or quite free. 
Petals naked; inflorescence of one or more distichous spikes or rarely 

simple and polystichous Tillandsia 

Petals each bearing 2 scales on the inner surface Vriesia 

Petals joined or closely agglutinated for most of their length; inflorescence 

always of polystichous-flowered spikes Guzmania 

Appendage of the seed apical, folded over at maturity; sepals distinctly asym- 
metric in most species; flowers polystichous Catopsis 

Ovary inferior; fruit baccate; seeds naked; leaves mostly serrate. 
Ovaries remaining distinct; inflorescence without a foliaceous coma. 
Inflorescences several, axillary, sessile, dense; petals naked, connate by their 

margins Greigia 

Inflorescence single, central, mostly scapose. 

Petals fleshy, 25-40 mm. long, attached to the filament-tube but with free 
margins; inflorescence sessile or scapose, densely lepidote. . .Bromelia 
Petals thin; filaments not joined in a tube; inflorescence always scapose. 
Sepals soft; petals 4-5 cm. long; pollen without pores, suldate . .Billbergia 

Sepals mucronate or pungent or the petals small; pollen with pores or 
aborted. 

Anthers bearing 2 large scales dorsally; petals naked; inflorescence 
compound; flowers sessile, subfasciculate or laxly polystichous. 

Androlepis 
Anthers naked. 

Inflorescence densely bipinnate with strobilate spikes; floral bracts 

entire; flowers compressed; epigynous tube lacking . Hohenbergia 

Inflorescence either simple or laxly compound or if densely bipinnate 

then the floral bracts serrate or the flowers distichous . .Aechmea 

Ovaries fusing to form a compound fruit; inflorescence bearing a coma of sterile 

foliaceous bracts at the apex Ananas 



AECHMEA R. & P. 

Stemless herbs; leaves rosulate; scape usually conspicuous; scape-bracts usu- 
ally bright-colored; inflorescence various; flowers sessile or rarely pedicellate; sepals 
usually asymmetric and mucronate; petals free, each bearing 2 scales or folds; 



382 FIELDIANA: BOTANY, VOLUME 24 

second series of stamens more or less adnate to the petals; pollen-grains with 
2 or 4 pores or sometimes aborted. 

About 150 species in tropical America. 

Flowers slenderly pedicellate. 

Sepals 6 mm. long; lower branches of the inflorescence much divided. 

Ae. mexicana 
Sepals 3.5 mm. long; lower branches of the inflorescence simple or slightly 

divided Ae. Luddemanniana 

Flowers sessile. 

Spikes polystichous-flowered. 

Inflorescence amply paniculate, elongate; spikes 4-8-flowered Ae. Iguana 

Inflorescence simple or densely digitate from a few spikes; spikes many- 
flowered. 

Floral bracts serrate, up to 65 mm. long Ae. magdalenae 

Floral bracts entire and short, or wanting. 

Sepals unarmed; floral bracts enfolding the ovary, thick . . Ae. bromeliifolia 

Sepals mucronate; floral bracts minute or wanting Ae. nudicaulis 

Spikes distichous-flowered. 

Floral bracts alate-decurrent and forming a pouch about the flowers, 10-17 

mm. long Ae. tillandsioides var. Kienastii 

Floral bracts not at all decurrent, 5-8 mm. long Ae. bracteala 

Aechmea bracteata (Sw.) Griseb. Fl. Brit. W. Ind. 592. 1864. 
Bromelia bracteata Sw. Prodr. 56. 1788. Ae. Schiedeana Schlecht. 
Linnaea 18: 437. 1844. Ae. laxiflora Benth. Bot. Voy. Sulph. 173. 
1846. Ae. regularis Baker, Jour. Bot. 17: 229. 1879. Ae. macracan- 
tha Brongn. ex Andre", 111. Hortic. 27: 59. 1880. Ae. Barleei Baker, 
Card. Chron. II. 20: 102. 1883. Ae. isabellina Baker, Jour. Bot. 28: 
305. 1890 (type from Boca del Polochic, Dept. Izabal, J. D. Smith 
1824). Izchu, Chuek (Pete"n, fide Bartlett); Ixchu, Tinajero (Pete"n, 
fide Lundell). 

Saxicolous or epiphytic in dense masses, forests, especially in 
rather arid regions, 50-900 meters; Pete"n; Alta Verapaz; Izabal. 
Mexico to Colombia. 

Plants 5-17 dm. or more high; leaves 3-13 dm. long, their sheaths elliptic, 
large; scape erect; scape-bracts lanceolate, entire, bright red; inflorescence laxly 
paniculate; spikes laxly 4-17-flowered, rhachis slender, flexuous; floral bracts 
broadly ovate, 5-8 mm. long, entire; flowers sessile; sepals asymmetric, 3-4 mm. 
long; petals 1 cm. long, yellow; ovary subglobose, enlarged in fruit. 

The berries are said to be edible. The hollow ellipsoid leaf- 
rosettes hold water which is useful to travelers in dry areas. The 
base is sometimes occupied by ants. In British Honduras the species 
is called "wild pine," but this name is applied to almost any con- 
spicuous bromeliad. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 383 

Aechmea bromeliifolia (Rudge) Baker in Benth. & Hook. Gen. 
PI. 3: 664. 1883. Tillandsia bromeliifolia Rudge, PI. Guian. 32. 1807. 

Rare epiphyte in forests; Pete"n (Uaxactun; Xaxha-Remate road). 
Not known in North America except in Guatemala and British Hon- 
duras; in South America from Colombia, Venezuela, and eastern 
Brazil to northern Argentina. 

Plant 7-9 dm. high; leaf-sheaths large, forming a tube, blades 4-9 cm. wide, 
laxly serrate; scape white-lanate, its bracts large, lance-ovate, entire, thin; inflo- 
rescence simple, strobilate, 15 cm. long, 3-4 cm. thick, densely white-lanate; floral 
bracts reniform, unarmed, bicarinate; sepals suborbicular, 7 mm. long, exceeding 
the floral bracts; petals 15 mm. long, soon black, bearing 2 fimbriate scales. 

Aechmea Iguana Wittm. Bot. Jahrb. 14: Beibl. 32: 3. 1891. 

Known only from the type, Bernoulli & Cario 695, from Quezal- 
tenango (Costa Cuca, between Cavallo Blanco and Ocos). 

Leaves unknown; scape-bracts laxly serrate; inflorescence amply paniculate; 
primary bracts lance-oblong, 15-18 cm. long, 40-45 mm. wide, serrate toward base; 
branches 10-12 cm. long; spikes short, 4-8-flowered; floral bracts triangular-ovate, 
mucronate, 10-12 mm. long; flowers polystichous, sessile; sepals lanceolate, 15 mm. 
long including the 5 mm. mucro; petals 15 mm. long, bearing 2 subdenticulate 
scales above the base; stamens included; ovary 15 mm. long, 8-10 mm. in diameter. 

I have seen nothing but the description of this species and am 
not even sure that it belongs in Aechmea. However, the descrip- 
tion is sufficient to indicate that the species is distinct from all other 
bromeliads known from Guatemala. 

Aechmea Luddemanniana (K. Koch) Brongn. ex Mez, Pflan- 
zenreich IV. 32: 120. 1934. Pironneava Luddemanniana K. Koch, 
Wochenschr. Gartn. 9: 182. 1866. Lamprococcus caerulescens Regel, 
Acta Hort. Petrop. 1: 93. 1871. Aechmea caerulescens Baker, Jour. 
Bot. 17: 227. 1879. 

Terrestrial or epiphytic in forest, 270-870 meters; Alta Verapaz 
(Finca Chama, valley of Rio Seniso, L. C. Stuart 24)- Vera Cruz; 
British Honduras. 

Plant 25-70 cm. high; leaves nearly straight, 3-6 dm. long, sheaths large, ellip- 
tic, blades ligulate, 4-7 cm. wide, spines 1-2 mm. long; scape erect, slender, scape- 
bracts imbricate, elliptic to linear-lanceolate, entire, membranaceous; inflorescence 
amply compound, cylindric to slenderly pyramidal, 12-30 cm. long, white-farinose; 
primary bracts narrow, mostly shorter than the branches; racemes laxly few-flow- 
ered; floral bracts filiform, usually shorter than the pedicels; pedicels slender, 6 mm. 
long, divergent; sepals asymmetric, mucronate, 3.5 mm. long; petals 9 mm. long, 
rose and blue; ovary 6 mm. long; berry 12 mm. long, bluish white. 

Called "wild pine" and "gallinasco" in British Honduras. 



384 FIELDIANA: BOTANY, VOLUME 24 

Aechmea magdalenae (Andre") Andre" ex Baker, Handb. Bromel. 
65. 1889. Chevalliera Magdalenae Andre", Enum. Bromel. 3. 13 Dec. 
1888; Rev. Hortic. 60: 563. 16 Dec. 1888. Bromelia Magdalenae 
C. H. Wright, Kew Bull. 1923: 267. 1923. Ananas magdalenae 
Standl. ex Standl. & Cald. Lista Prelim. PI. S. Salvador 45. 1925. 
Silkgrass; Pita floja (Alta Verapaz, Central America, fide Standley) ; 
Pita (Pete*!!, fide Bartlett); Pinuela (Pete*n, fide Standley); Man 
(Quecchi). Figure 65. 

Terrestrial in forests and thickets, usually below 500 meters; 
Pete"n (Tikal). Mexico to Ecuador, the species described from the 
Rio Magdalena in Colombia. 

Plant 10-15 dm. high; leaves coarse, up to 2 meters long, blades linear, 5-10 
cm. wide, often bright red (Standley), laxly serrate with curved teeth 5 mm. long; 
scape stout, its bracts foliaceous, the upper ones massed below the inflorescence 
and reflexed; inflorescence of a few sessile globose spikes in a dense mass or rarely 
simple; floral bracts recurving, acuminate, 65 mm. long, densely serrate, thick, 
coriaceous; flowers sessile, 5 cm. long; sepals triangular, unequal, 35-38 mm. long; 
petals 4 cm. long; ovary elliptic, complanate. 

According to Standley this species is common in the North Coast 
(Izabal) and lowlands of Baja Verapaz, apparently also in lowlands 
of northern Huehuetenango. Large amounts of the fiber are said to 
be separated in northern Huehuetenango. It is cultivated occasion- 
ally for ornament or as a curiosity in Guatemala City and probably 
elsewhere. The leaves are retted in water, and fiber is usually ex- 
tracted by pounding them on stones in running water. It is a very 
fine and tough fiber, much used for hammocks, bags and string. 

Aechmea mexicana Baker, Jour. Bot. 17: 165. 1879. Ae. 
Bernoulliana Wittm. Bot. Jahrb. 14: Beibl. 32: 1. 1891. 

Epiphytic in forests, 250-1300 meters; San Marcos. Vera Cruz; 
Costa Rica; Ecuador. 

Plants often over a meter high; leaves many in a utriculate rosette, blades ligu- 
late, 6-12 cm. wide, serrate; scape stout; scape-bracts narrowly lanceolate, entire, 
stramineous; inflorescence amply paniculate, furfuraceous; racemes laxly few-flow- 
ered, floral bracts much shorter than the pedicels, filiform; pedicels 4-16 mm. long; 
sepals strongly asymmetric, mucronate, 6 mm. long; petals 10-15 mm. long, red or 
lilac; ovary 6 mm. long, often enlarging in fruit. 

Aechmea nudicaulis (L.) Griseb. Fl. Brit. W. Ind. 593. 1864. 
Bromelia nudicaulis L. Sp. PI. 286. 1753. ?Tillandsia serrata Sesse" 
& Moc. Fl. Mex. ed. 2. 81. 1894. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 385 




FIG. 65. Aechmea magdalenae. Inflorescence and upper part of scape (X 2 A). 



Epiphytic in forests, 20-1050 meters; southern Mexico to Pan- 
ama; West Indies; a variety in South America; probably extending 
into Guatemala although not recorded yet. 

Plant very variable in proportions, 3-7 dm. high; leaves densely fasciculate, 
3-10 dm. long, sheaths large, elliptic, dark, forming an urceolate pseudobulb, blades 
ligulate, broadly obtuse and apiculate, 6-10 cm. wide, densely pale-lepidote be- 
neath, armed with coarse black teeth to 4 mm. long; scape slender, white-floccose; 
scape-bracts imbricate, congested below the inflorescence, elliptic, entire, red; in- 



386 



FIELDIANA: BOTANY, VOLUME 24 



florescence spicate, fertile throughout, 5-25 cm. long; floral bracts small or some- 
times wanting, entire; flowers 22 mm. long; sepals free, very asymmetric, mucro- 
nate, 5-10 mm. long; petals 12 mm. long, yellow, bearing 2 fimbriate scales; ovary 
subglobose. 




FIG. 66. Aechmea tillandsioides var. Kienastii. Portion of scape and inflo- 
rescence and apex of leaf (X K)- 

Aechmea tillandsioides (Mart.) Baker var. Kienastii (E. 
Morr. ex Mez) L. B. Smith, Caldasia 5: 5. 1942. Ae. Kienastii 
E. Morr. ex Mez in DC. Monog. Phan. 9: 243. 1896. Ae. squarrosa 
Baker, Jour. Bot. 28: 305. 1890 (type from Boca de Polochic, Dept. 
Izabal, J. D. Smith 1823), not Baker, 1889. Pie de gallo (Pete'n); 
eck (Alta Verapaz). Figure 66. 

Epiphytic in forests, up to 300 meters; Pete'n; Alta Verapaz; 
Izabal. Southern Mexico; Central America; Colombia; Amazon 
Basin. The typical variety is in Colombia, the Amazon Basin and 
Guiana. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 387 

Leaves 5-9 dm. long with serrate blades 10-65 mm. wide; scape short, its 
bracts remote, lanceolate, red, serrate; inflorescence simple or digitate (pinnate in 
the typical variety), white-floccose at first; spikes oblong, 4-11 cm. long; rhachis 
excavated; floral bracts distichous, imbricate, 10-17 mm. long, convex, nerved; 
sepals asymmetric, 7-10 mm. long; petals dark purple. 



ANANAS Mill. 

Plant stemless, not stoloniferous; leaves densely rosulate, scarcely enlarged at 
base; scape evident, erect; inflorescence densely strobiliform, crowned with a tuft 
of sterile foliaceous bracts, frequently producing slips at its base; flowers sessile; 
sepals free, obtuse, slightly asymmetric; petals free, erect, violet or red, each bear- 
ing 2 slenderly infundibuliform scales; stamens included; pollen-grains ellipsoid 
with 2 pores; ovaries coalescing with each other and with the bracts and axis to 
form a fleshy compound fruit, epigynous tube short; ovules borne near top of cell. 

At least four species and numerous cultivated forms. The genus 
is a native of Brazil and Paraguay but has been so widely cultivated 
even before the discovery of America that its natural limits are dif- 
ficult to define. 

Ananas comosus (L.) Merrill, Interpr. Rumph. Amboin. 133. 
1917. Bromelia Ananas L. Sp. PL 285. 1753. B. comosa L. Herb. 
Amboin. 21. 1754. Ananassa saliva Lindl. Bot. Reg. 13: under pi. 
1068, nomen nudum. 1827. Ananas sativus Schultes in R. & S. Syst. 
Veg. 7: 1283. 1830. A. Ananas Voss. in Vilm. Blumeng. ed. 3. 1: 964. 
1895. Pina; Pine; Pineapple; Chop (Quecchi, Poconchi, Cacchi- 
quel). Figure 67. 

Leaves coarsely and laxly spinose-serrate; scape short, stout; scape-bracts ser- 
rate; inflorescence large, many-flowered; floral bracts soon exposing the tops of the 
ovaries, relatively inconspicuous, weakly serrulate or entire; syncarp well over 
15 cm. long at maturity with copious palatable flesh; seeds lacking or very rare. 

Native of southern Brazil and Paraguay, but now cultivated in 
most tropical regions throughout the world. Standley notes that the 
date of its introduction into Guatemala is unknown, but probably it 
was brought up from South America along the Atlantic coast by the 
Caribs, like yuca (mandioca). Grown commonly in Guatemala from 
about 1500 meters down to sea level, but much more common at low 
elevations, especially along the Pacific plains. Large amounts of 
fruit are carried up into the higher mountains for sale in the markets. 
Ordinarily the fruits sell at 3-6 cents each, but about Coban fairly 
good ones are sometimes offered as low as one cent. The plants have 
run wild in some places, especially about Puerto Barrios (Izabal), 
but the wild plants do not produce fruits fit to eat as a rule. The 
sprout shoots, while young and relatively tender, are cooked and 




FIG. 67. Ananas comosus. a, Inflorescence and leaf (X K). b, Floral bract 
with ovary (X 1). c, Floral bract with ovary, side view (XI). 



388 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 389 

eaten commonly. They are called "hijos de pina." Bunches of such 
sprouts often are offered for sale in the markets of the Oriente, par- 
ticularly at Zacapa and Jutiapa. 

ANDROLEPIS Brongn. ex Houllet 

Large showy herbs; leaves rosulate, spinose-serrate; scape central, erect; inflo- 
rescence compound with simple reduced or divided branches; spikes polystichous- 
flowered; flowers sessile, perfect; sepals free, asymmetric, mucronate; petals short- 
connate, naked; stamens included, filaments short, adnate; anthers bearing 2 foli- 
aceous appendages at apex; ovary wholly inferior, epigynous tube short; ovules 
numerous, borne along the upper part of the cell, acute. 

The genus consists of the two species noted here. 

Branches of the inflorescence well developed and the lower ones often divided. 

A. Donnell-Smithii 
Branches of the inflorescence very short; flowers subfasciculate A. Skinneri 

Androlepis Donnell-Smithii (Baker) Mez in DC. Monog. 
Phan. 9: 161. 1896. Aechmea Donnell-Smithii Baker, Jour. Bot. 28: 
305. 1890. 

Terrestrial in coastal woods, sea level to 75 meters; Izabal (type 
from Livingston, Rio Dulce, J. D. Smith 1825}. Honduras; Costa 
Rica. 

Plant over 1 meter high; leaves erect, stiff, 0.5 to over 1 meter long, sheaths 
elliptic, entire, blades ligulate, acute or acuminate, 4-8 cm. wide, flat with involute- 
pungent apex, densely serrate with spines to 3 mm. long, glabrous above, finely 
pale-lepidote between the nerves beneath; scape 1 cm. or thicker, buff-furfura- 
ceous; scape-bracts lanceolate, acute, densely imbricate, membranaceous toward 
base, thick coriaceous toward apex, pungent, pale-lepidote; inflorescence amply 
compound, usually dense, narrowly pyramidal or cylindric, furfuraceous; primary 
bracts linear, membranaceous, deflexed, the upper much reduced; branches spread- 
ing, well developed, the lowest usually divided; spikes usually lax, longer than 
broad, 4-5-flowered; floral bracts minute, shorter than the ovary, acuminate, pun- 
gent; flowers suberect, to 14 mm. long; sepals broadly triangular with a large wing, 
5-6 mm. long, coriaceous, soon glabrous; petals elliptic, 1 cm. long; stamens in- 
cluded; ovary usually not much enlarged in fruit. 

Steyermark notes that this species grows just inside the border 
of the woods above the beach. Called "gallinazo" in Honduras. 

Androlepis Skinneri (K. Koch) Brongn. ex Houllet, Rev. 
Hortic. 42: 12. 1870. Pothuava Skinneri K. Koch, Wochenschr. 
Gartn. 4: 190. 1861 (based on plants of unknown origin cultivated 
in Germany). Billbergia Skinneri K. Koch, I.e. Aechmea leuco- 
stachys Baker, Handb. Bromel. 39. 1889. Ae. Skinneri Baker, op. 
cit. 49. Figure 68. 




FIG. 68. Androlepis Skinneri. a, Habit (X 1/5). b, Appendaged anther, 
ventral view (X 8). c, Appendaged anther, dorsal view (X 8). d, Longitudinal 
section of flower (X 2). 



390 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 391 

On rocks or epiphytic in forests, usually forming large masses on 
trunks or in crotches of trees, at low altitudes; Pete"n (La Libertad, 
Lundell 2605, 2986} . British Honduras. 

Leaves 6 dm. long, sheaths large, suborbicular, blades ligulate, 5-8 cm. wide, 
finely pale-lepidote beneath, densely serrulate; scape 3 dm. long, stout; scape- 
bracts imbricate, acute, serrulate, stramineous; inflorescence lax, subcylindric, 
2 dm. long; primary bracts narrowly triangular, entire; spikes suberect, 1-5-flow- 
ered; floral bracts minute or wanting; sepals 5 mm. long; petals elliptic, 1 cm. long, 
flavous; ovary stout, much enlarged in fruit. 

BILLBERGIA Thunb. 

Plants stemless; leaves rosulate or rarely fasciculate, sheaths large, blades 
mostly ligulate, spinose-serrate, often banded; scape erect or arching; scape-bracts 
red, membranaceous; inflorescence simple or compound; flowers large, showy, ses- 
sile or pedicellate; sepals free, erect; petals free, bearing 2 scales on the inner surface 
near the base, the claw long, the blade narrow; stamens exserted at anthesis, both 
series free or the second adnate to the petals up to the scales, pollen-grains with 
longitudinal folds when dry but no pores; style exceeding the stamens, epigynous 
tube large, ovules many. 

About 50 species, the great majority of which are native to eastern 
Brazil; the remainder are in Mexico, Central America, Trinidad, and 
Venezuela and Colombia to northern Argentina. 

Flowers sessile; floral bracts suborbicular B. pallidiflora 

Flowers slenderly long-pedicellate; floral bracts narrowly triangular. .B. viridiflora 

Billbergia pallidiflora Liebm. Ind. Sem. Hort. Haun. 1854: 26. 
1854; Ann. Sci. Nat. IV. 2: 373. 1854. 

Oak-pine woods, 1200-1500 meters; Zacapa (along upper reaches 
of Rio Sitio Nuevo, between Santa Rosalia and first waterfall, Steyer- 
mark 42264). Mexico; Nicaragua. 

Leaves tubular-rosulate, 6 dm. long, blades ligulate, acute, 45 mm. wide, spi- 
nose-serrate; scape arching; scape-bracts densely imbricate, lance-elliptic, over 
15 cm. long; inflorescence simple, densely cylindric, many-flowered, 2 dm. long, 
densely white-farinose; floral bracts much shorter than the ovary; sepals ovate- 
oblong, broadly acute and apiculate, unequal, 9-12 mm. long; petals linear, acute, 
to 52 mm. long, green, imperfectly revolute; ovary slenderly ovoid, 12-18 mm. 
long, coarsely sulcate with the ridges soon glabrous, epigynous tube narrow, 
6 mm. long. 

Billbergia viridiflora H. Wendl. Allg. Gartenz. 22: 154. 1854. 
Figure 69. 

On rocks and epiphytic in forests, at low elevations; Pete"n; Alta 
Verapaz; Izabal. Tabasco; British Honduras. 



392 



FIELDIANA: BOTANY, VOLUME 24 




FIG. 69. Billbergia viridiflora. Portion of scape and inflorescence (X K). 

Leaves 12-15 in a tubular rosette, green, minutely white-lepidote, sheaths 
narrow, entire, blades ligulate, acuminate, 4-9 cm. wide, teeth 1-4 mm. long; 
scape suberect to arching, slender, about equaling the leaves; scape-bracts imbri- 
cate, up to 14 cm. long, serrulate; inflorescence simple, lax, 2-5 dm. long; floral 
bracts narrow, to 2 cm. long; pedicels spreading, slender, up to 5 cm. long; flowers 
glabrous; sepals acuminate, 21 mm. long, green; petals 4-5 cm. long, acute, green, 
bearing 2 subentire scales at base; ovary ellipsoid, green. 



BROMELIA L. 

Coarse terrestrial herbs, spreading by rhizomes; leaves usually rosulate, with 
large curved spines along the margin; inflorescence sessile or stipitate, always com- 
pound; sepals free or somewhat united, obtuse or acute, rarely mucronulate; petals 
rarely with a definite claw, centrally united by the filaments but their margins free, 
naked, fleshy in most species; stamens included, the filaments forming a tube for 
part of their length, anthers narrow, acute; ovary passing gradually into the thick 
pedicel, epigynous tube conspicuous to nearly lacking; berry succulent, relatively 
large; seeds few to many, flattened. 

Nearly 40 species in tropical America. 

Inflorescence an elongate lax panicle, terminating a definite scape. 

Floral bracts and sepals subulate; petals densely white-tomentose toward apex. 

B. Pinguin 

Floral bracts and sepals triangular; petals sparsely lepidote to glabrous toward 
apex B. sylvestris 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 393 

Inflorescence a densely capituliform panicle, scapeless and sunk in the center of the 

leaf-rosette. 
Filament- tube over 2 cm. long; indument of very coarse black-brown scales. 

B. Karatas 
Filament-tube not over 12 mm. long; indument of fine buff or white scales. 

B. Wercklei 

Bromelia Karatas L. Sp. PI. 285. 1753. Karatas Plumieri E. 
Morr. Belg. Hortic. 22: 131. 1872. Cham, Chom (Pet&i, Maya); 
Pinuela (Pete"n, Huehuetenango) ; Pina (Alta Verapaz) . Figure 70, 
c, d. 

In marginal forests bordering savannas and in deciduous forests, 
1300-1500 meters; Pete"n; Alta Verapaz; Sacatepe"quez; Huehuete- 
nango. Mexico and the West Indies to Ecuador and Brazil. 

Rosette up to 3 meters in diameter, leaf-sheaths large, covered with long coarse 
dark brown scales, blades 3-5 cm. wide, minutely lepidote beneath, teeth coarse, 
5-8 mm. long; inflorescence a many-flowered capituliform panicle, flat-topped, 
surrounded by the red inner leaves; primary bracts foliaceous; floral bracts nar- 
row, oblanceolate, thin, coarsely lepidote; flowers 6-9 cm. long; pedicels short, 
stout; sepals lanceolate, acute, lepidote; petals 4 cm. long, glabrous, rose with 
white base and margins; ovary lepidote; berry fusiform, 8 cm. long, 2 cm. thick, 
acid, edible. 

According to Standley the species is much planted in hedges in 
Guatemala, and is common about Antigua (Sacatepe"quez, 1500 me- 
ters) and other places of lower elevation. Sometimes called "pinuela 
de cabeza de negro" in Guatemala. Flowering and fruiting in the 
rainy season. The handsome heads of flowers and fruits are often 
sold in the markets for use in decorating nacimientos and altars. In 
El Salvador the ripe fruits of this species or the closely allied B. 
Wercklei are cooked with sugar and used to make the beverage Atol 
de pina or Atol de pinuela. The young shoots are called pollas there; 
they are cooked in soup, fried with eggs, and used in other ways for 
food. The young inflorescences are known there as motates. 

Bromelia Pinguin L. Sp. PL 285. 1753. Pinuela; Ixchuu 
(Pet^n, Maya). Figure 70, a. 

Common and widely distributed in the lowlands of Guatemala, 
mostly or wholly at 1100 meters or lower. Known definitely (accord- 
ing to field and other records) from Pete'n; Baja Verapaz; Zacapa; 
Chiquimula; El Progreso; Guatemala; Escuintla; Jutiapa; Quezalte- 
nango; probably in all the lowland departments. 

Plant 1 meter high; leaves many, over 2 meters long, sheaths broad, tomentose- 
lepidote, blades 4 cm. wide, deep green above, pale-lepidote beneath, teeth 10 mm. 



394 



FIELDIANA: BOTANY, VOLUME 24 



long; scape stout, white-floccose; scape-bracts subfoliaceous, reddish; inflorescence 
narrow, white-floccose; primary bracts like the scape-bracts; floral bracts linear- 
subulate, 3 cm. long; flowers 6 cm. long; sepals triangular-subulate, pale; petals 
narrow, 3 cm. long, rose with white base and margins, densely white-tomentose 
toward apex; berry yellow, very acidulous, aromatic, edible. 




FIG. 70. a, Bromelia Pinguin, branch of inflorescence ( X Yz). b, B. sylvestris, 
branch of inflorescence and primary bract (X M)- c > d, B. Karatas: c, flowers 
(X M); d, corolla cut open (X %). 



Known in Yucatan by the Maya names "tsalbay," "cham," 
"chom" and "hman." In Guatemala young shoots are called "hijos 
de pina." The plant often forms wide and dense thickets on the 
plains of the Pacific coast and all through the lower Motagua Valley. 
It is much planted for hedges, even in regions where it is not native. 
The young inflorescences are called "mutas" in Guatemala. The 
"hijos" are much used as food in all the dryer regions. They can be 
had when ordinary vegetables are scarce or unobtainable, as at the 
end of the dry season. The fruit is intensely sour and acrid. It is 
sometimes used for making vinegar. 

Bromelia sylvestris Willd. ex Link, Enum. 1: 308. 1821. 
Figure 70, 6. 

Arid woodlands, at low elevations; Vera Cruz; Yucatan; Hon- 
duras; probably extending into Guatemala. 

Flowering plant nearly 1 meter high; leaves many, about 1 meter long, sheaths 
large, white-lepidote, blades glabrous and lustrous above, densely pale-lepidote 
beneath, teeth 5 mm. long; scape stout, white-floccose; scape-bracts subfoliaceous, 
reddish; inflorescence narrow, white-floccose; primary bracts like the scape-bracts; 
floral bracts narrowly triangular, flat, thin; flowers 5 cm. long; sepals lance-triangu- 
lar, acute or obtuse, 15-18 mm. long; petals lanceolate, obtuse, 25 mm. long, blue 
or rose with white margins, glabrous or sparsely lepidote at apex, connate for only 
3 mm. with the filament- tube. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 395 

Some of the supposed records of the closely allied B. Pinguin 
may represent this species. 

Bromelia Wercklei Mez, Repert. Sp. Nov. 16: 2. 1919. 

From 40 to 1500 meters; Mexico; El Salvador; Nicaragua; Costa 
Rica; probably extending into Guatemala. 

Leaves 1-2 meters long, sheaths densely pale-tomentose-lepidote on both sides, 
blades linear, 3-4 cm. wide, acuminate, laxly serrate with teeth 7 mm. long; inflo- 
rescence many-flowered, dense, sunk in the center of the leaf-rosette, densely lepi- 
dote with fine elongate buff scales; primary bracts large with very broadly ovate 
base and long foliaceous blade; floral bracts narrowly oblanceolate, 45 mm. long, 
the upper half coarsely laciniate-serrate; flowers to 6 cm. long; sepals oblong, entire, 
acute, 24 mm. long; petals ligulate, acute, 3 cm. long, connate for 12 mm. with the 
filament-tube, fleshy, glabrous or slightly puberulous toward the apex; ovary 
25 mm. long. 

Cultivated as a hedge. The acid fruits are used in soft drinks. 
Called "piiiuela casera" in Costa Rica, "piiia de cerco" in El Salvador 
and "piriuela" and "timbiriche" in Mexico. It is not always certain 
whether records apply to this species or to the closely allied and 
better known B. Karatas. 

CATOPSIS Griseb. 

Stemless herbs; leaves utriculate-rosulate with large sheaths, minutely lepi- 
dote, green; scape evident; inflorescence usually compound; flowers polystichous, 
usually sessile, perfect or functionally dioecious; sepals free, glabrous, usually 
asymmetric; petals free, naked; stamens included; ovary superior, style very short 
or none; seeds with an apical folded coma. 

Twenty-five species in Florida, Mexico, Central America, the 
West Indies and northern South America. 

Flowers slenderly pedicellate C. pedicellata 

Flowers sessile or subsessile. 
Sepals 10-18 mm. long. 

Floral bracts exceeding the sepals. 
Floral bracts cucullate; inflorescence pendulous or erect; flowers perfect. 

C. cucullata 
Floral bracts straight toward apex; inflorescence erect; flowers dioecious. 

C. triticea 
Floral bracts shorter than the sepals, straight toward apex. 

Sepals subsymmetric, 15-18 mm. long C. Wawranea 

Sepals distinctly asymmetric, 10-15 mm. long. 

Petals bright yellow, much exceeding the submembranaceous sepals; 

scape slender, usually decurved C. nutans 

Petals white, equaling or shorter than the coriaceous sepals; scape erect, 
stout . . . . C. Berteroniana 



396 FIELDIANA: BOTANY, VOLUME 24 

Sepals not more than 9 mm. long. 

Scape-bracts all shorter than the internodes. 

Sepals not over 4.5 mm. long; inflorescence delicate with very slender axes; 

leaves mostly 10-15 cm. long C. apicroides 

Sepals 5-9 mm. long. 

Leaves in a cylindric rosette, usually strict; sepals cuneate on the right 
with the wing much exceeding the midnerve, 5-6 mm. long. 

C. nitida 

Leaves in a crateriform rosette, arching-divergent; sepals curved on both 
sides of the base with the wing scarcely if at all exceeding the mid- 
nerve, 7-8 mm. long. 

Flowers perfect, suberect; capsule short-beaked C. sessiliflora 

Flowers dioecious, spreading; capsule beakless C. aloides 

Scape-bracts equaling or exceeding the internodes, or rarely the uppermost 
slightly shorter. 

Inflorescence pendulous, 3-5 dm. long C. pendula 

Inflorescence erect or somewhat curved but never pendulous. 

Floral bracts equaling the sepals ,. C. triticea 

Floral bracts distinctly shorter than the sepals at maturity. 

Leaves with conspicuous pale or white margins, 10-18 cm. long. 

Leaf-blades narrowly triangular, acuminate C. brevifolia 

Leaf-blades ligulate, broadly acute or obtuse, apiculate. 

C. Morreniana 
Leaves concolorous. 

Leaf-blades 10-45 mm. wide. 
Spikes dense or subdense. 

Lower primary bracts more than half as long as the axillary 
branches, usually exceeding them, ample, bladeless. 

C. Hahnii 

Lower primary bracts much shorter than the axillary branches, 
laminate. 

Spikes short-stipitate, spreading, very dense C. subulata 

Spikes long-stipitate, strict, subdense C. floribunda 

Spikes lax. 

Branches of the inflorescence strict; spikes long-stipitate. 

C. floribunda 
Branches of the inflorescence divergent to spreading; spikes 

short-stipitate. 
Lower primary bracts nearly equaling the usually simple 

branches C. delicatula 

Lower primary bracts mostly shorter than the sterile bases 

of the usually divided branches C. montana 

Leaf-blades not over 5 mm. wide; primary bracts much shorter than 
the sterile bases of the spikes. 

Leaf-sheaths conspicuous, large C. juncifolia 

Leaf-sheaths inconspicuous, narrow C, Lundelliana 

Gatopsis aloides (Schlecht. & Cham.) Baker, Handb. Bromel. 
154. 1889. Tillandsia aloides Schlecht. & Cham. Linnaea 6: 55. 1831. 

Epiphytic in forest, 100-600 meters; Pete'n; Alta Verapaz; 
Huehuetenango. Southern Mexico; British Honduras. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 397 

Plant 14-46 cm. high; leaves few in a slenderly crateriform rosette, to 25 cm. 
long, sheaths inconspicuous, about as long as the blades, blades ligulate, broadly 
acute or obtuse, apiculate, 23 mm. wide; scape erect or arching, slender, angled or 
narrowly winged; scape-bracts erect, remote, broadly elliptic-ovate; inflorescence 
simple or laxly compound from 2-4 spikes, to 15 cm. long, glabrous; primary bracts 
like the scape-bracts, usually shorter than the sterile bases of the branches; spikes 
dense except toward base, 3-13 cm. long; floral bracts ovate, obtuse, much shorter 
than the sepals, thin, nerved; flowers spreading, dioecious; sepals asymmetric, 
broadly subelliptic, 8 mm. long; petals slightly exserted, yellow; capsule ovoid, 
acute but not beaked, 10-15 mm. long. 

Catopsis apicroides (Schlecht. & Cham.) Baker, Jour. Bot. 25: 
174. 1887. Tillandsia apicroides Schlecht. & Cham. Linnaea 6: 55. 
1831. Figure 71. 

Epiphytic in forests, 1-1140 meters; Izabal. Southern Mexico; 
Central America. 

Plants 12-45 cm. long; leaves few, their blades ligulate, 10-25 mm. wide, the 
inner rounded-apiculate; scape more or less curved, very slender; scape-bracts 
remote; inflorescence compound, lax; primary bracts very short; spikes many- 
flowered, 2-11 cm. long; floral bracts equaling or shorter than the sepals; flowers 
dioecious; sepals asymmetric, 4.5 mm. long; petals 6 mm. long, yellow. 

Catopsis Berteroniana (Schultes) Mez in DC. Monog. Phan. 
9: 621. 1896. Tillandsia Berteroniana Schultes in R. & S. Syst. 7: 
1221. 1830. 

Epiphytic in thickets and forests, at low altitudes; Pete"n (La Lib- 
ertad, Lundell 2909}. Southern Florida; Bahamas; Cuba; Jamaica; 
Santo Domingo; British Honduras; Panama; Trinidad; British Gui- 
ana; eastern Brazil. 

Plant 4-9 dm. high; leaves to 4 dm. long, white-cretaceous, their sheaths very 
large, blades triangular, 4-5 cm. wide; scape erect, stout; lower scape-bracts imbri- 
cate, foliaceous, the upper ovate, often remote; inflorescence usually compound 
with short, broadly ovate primary bracts; spikes long-stipitate, laxly flowered; 
floral bracts broad, obtuse, 6-8 mm. long; flowers perfect, suberect; sepals asym- 
metric, obovate, 10-12 mm. long, equaling or exceeding the white petals. 

Catopsis brevifolia Mez & Werckle" ex Mez, Bull. Herb. Boiss. 
II. 4: 1127. 1904. 

Epiphytic; Suchitepe"quez (Volcan Santa Clara, 1250-2650 me- 
ters, Steyermark 46648). Costa Rica. 

Plant 2-3 dm. high; leaves many in a dense rosette, with or without a creta- 
ceous coat, sheaths large, ovate or elliptic, blades narrowly triangular, acuminate, 



398 FIELDIANA: BOTANY, VOLUME 24 




FIG. 71. Catopsis apicroides. Habit (X >0. 

2 cm. wide, the margin pale; scape erect, slender; scape-bracts suberect, foliaceous, 
exceeding the internodes; inflorescence laxly and sparsely bipinnate (or the lowest 
branch divided again) ; lower primary bracts like the scape-bracts, longer or shorter 
than the branches, the upper much shorter; spikes subdense, distinctly stipitate; 
floral bracts broadly ovate, obtuse, shorter than the sepals, nerved; flowers sub- 
erect, dioecious, 6-7 mm. long; sepals strongly asymmetric; petals broadly ligulate, 
slightly longer than the sepals; stamens unequal; ovary pyramidal-ovoid; stigmas 
subsessile. 

I have not seen any material of the type and the immature Gua- 
temalan material is associated with it solely on the basis of the type 
description. 

Catopsis cucullata L. B. Smith, Contr. Gray Herb. 104: 72. 
1934. Chopixul (fide Standley). Figure 72. 

Epiphytic, 1290 meters; Alta Verapaz (Coban, Johnson 569). 
Jalapa; Vera Cruz. 

Leaves up to 22 cm. long, thin, their sheaths indistinct, blades subtriangular, 
acute and apiculate, 2 cm. wide; scape erect or decurved; scape-bracts longer than 
the internodes, ovate, acute; inflorescence simple or few-branched; primary bracts 
shorter than the spikes; spikes suberect, to 10 cm. long; floral bracts ovate, acute, 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 399 




FIG. 72. Catopsis cucullata. a, Habit (X 
stamens shown (X 1). c, Pistil (X 1). 



). b, Single petal with three 



10-15 mm. long, exceeding the flowers, cucullate, thin; flowers perfect, fragrant 
(fide H. Johnson); sepals asymmetric, obovate, 10 mm. long; petals barely exceed- 
ing the sepals. 

Catopsis delicatula L. B. Smith, Contr. Gray Herb. 117: 4. 
1937. 

Epiphytic in forest, 100-670 meters; endemic; Suchitepe"quez; 
Retalhuleu (type from San Felipe, 670 meters, Rojas 445). 

Immature plant 28 cm. high; leaves few, subfasciculate, their sheaths small 
and indistinct, blades linear-triangular, acuminate, 12 mm. wide, flat; scape erect, 
very slender, its bracts erect, narrowly lanceolate, much exceeding the internodes; 
inflorescence laxly paniculate, axis geniculate; lower primary bracts about equal- 
ing the young spikes, the upper much reduced; spikes spreading, 4 cm. long, laxly 
few-flowered ; floral bracts ovate, shorter than the sepals of the more mature flow- 
ers; flowers spreading; sepals strongly asymmetric, 5 mm. long; petals 6 mm. long. 

The very immature state of the known collections of this 
species make it very difficult to judge what the relative size of parts 
would be in mature plants. However, the almost filiform tips of the 
scape-bracts and lower primary bracts are very distinctive. 

Catopsis floribunda (Brongn.) L. B. Smith, Contr. Gray Herb. 
117: 5. 1937. Pogospermum floribundum Brongn. Ann. Sci. Nat. V. 
1: 329, nomen provisorium. 1864. Catopsis nutans Griseb. as to de- 
scription and use by later authors, not as to basionym. Figure 73. 



400 



FIELDIANA: BOTANY, VOLUME 24 




FIG. 73. Catopsis floribunda. a, Inflorescence and top of scape (X 1). b, 
Sepal (X 5). c, Floral bract (X 20). d, Apex of leaf (X 1). 

Epiphytic in forests, up to 1500 meters; Baja Verapaz; Que- 
zaltenango. Florida; West Indies; British Honduras; Costa Rica; 
Venezuela. 

Plant 4-7 dm. high; leaves many, strict, 2-4 dm. long, their sheaths large, 
elliptic, blades narrowly triangular, 15-30 mm. wide; scape erect, slender, its 
bracts imbricate with the lower foliaceous; inflorescence laxly compound, 15-40 cm. 
long; primary bracts usually shorter than the sterile bases of the branches; spikes 
subdense or lax; floral bracts ovate, shorter than the sepals; flowers suberect, per- 
fect; sepals 4-6 mm. long, asymmetric; petals 7 mm. long, white. 

Catopsis Hahnii Baker, Jour. Bot. 25: 175. 1887. C. Oerstediana 
Mez in DC. Monog. Phan. 9: 630. 1896. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 401 

Epiphytic in forests, 1300-2500 meters; Alta Verapaz; Chimal- 
tenango; Solola; Quiche"; Huehuetenango; San Marcos. Southern 
Mexico; Honduras; Nicaragua. 

Plant to over 5 dm. high; leaves in a dense subfasciculate rosette, 2-4 dm. long, 
sometimes white-cretaceous beneath, sheaths elliptic, as long as or longer than the 
blades, blades triangular, acute or acuminate, 30-45 mm. wide; scape erect or arch- 
ing; scape-bracts foliaceous, large, densely imbricate; inflorescence densely bipin- 
nate or tripinnate, subthyrsoid, 11-25 cm. long, the staminate with more and 
smaller flowers; primary bracts suberect, lance-ovate, acuminate, usually exceed- 
ing the lower branches; spikes dense; floral bracts ovate-elliptic, obtuse, shorter 
than the sepals; flowers Suberect; sepals asymmetric, pistillate 8-9 mm. long, 
staminate 6 mm.; petals suborbicular, white, barely exceeding the sepals. 

Catopsis juncifolia Mez & Werckle" ex Mez, Bull. Herb. Boiss. 
II. 4: 1124. 1904. 

Epiphytic, between 150 and 700 meters; Alta Verapaz (Cerro 
Chinaja, Steyermark 45689). Costa Rica. 

Plant 23-27 cm. high; leaves many in a subbulbous rosette, 9-14 cm. long, 
sheaths large, ovate-elliptic, densely brown-punctulate-lepidote, blades junciform- 
subulate, acuminate, 5 mm. wide; scape erect, slender; at least the lower scape- 
bracts exceeding the internodes; inflorescence sparsely bipinnate, lax, strict, 
composed of a terminal spike of about 10 flowers and 2 long-stipitate few-flowered 
lateral spikes; primary bracts lance-ovate, acuminate, much shorter than the 
sterile bases of the spikes; floral bracts broadly ovate, obtuse, much shorter than 
the sepals; flowers suberect to spreading, dioecious, only the pistillate known; 
sepals asymmetric, 4-5 mm. long; petals barely exserted; ovary pyramidal. 

Gatopsis Lundelliana L. B. Smith, Contr. Gray Herb. 117: 6. 
1937. Figure 74. 

Epiphytic in forests, at low elevations; British Honduras (type 
from Valentin, El Cayo District, Lundell 6256); Panama; probably 
extending into Guatemala. 

Plant 25-30 cm. high; leaves many in a dense subglobose rosette, 1 dm. long, 
densely and obscurely punctulate-lepidote, sheaths narrowly ovate or elliptic, 15- 
30 mm. long, not inflated, blades linear, acuminate, 5 mm. wide, involute toward 
apex; scape erect, very slender, glabrous; scape-bracts exceeding the internodes, 
lanceolate with a long narrow acuminate blade; inflorescence lax, of 3-4 branches, 
7-12 cm. long; primary bracts like the upper scape-bracts, shorter than the sterile 
bases of the branches; spikes divergent, straight, very laxly subtristichous-flow- 
ered, to 95 mm. long; floral bracts broadly ovate, obtuse, thin, prominently nerved, 
much shorter than the sepals; flowers suberect; sepals obovate, obtuse, strongly 
asymmetric, 5 mm. long; petals barely exserted; style short but distinct. 

Catopsis montana L. B. Smith, Contr. Gray Herb. 117: 6. 1937. 
Epiphytic in oak woods, 1300-1700 meters; Jalapa; Solola. Cuba. 



402 



FIELDIANA: BOTANY, VOLUME 24 




FIG. 74. Catopsis Lundelliana. 
(X %). 6, Scape-bract (X %). 



a, Inflorescence and upper part of scape 



Plant 5-6 dm. high; leaves many in a cyathiform rosette, 35-45 cm. long, 
sheaths elliptic, about as long as the blades but indistinct, blades linear-lanceolate, 
acuminate, to 33 mm. wide; scape erect, slender; scape-bracts erect, the lower sub- 
foliaceous, imbricate, the upper ovate, acuminate, at least equaling the internodes; 
inflorescence laxly paniculate with the lower branches divided, 25-30 cm. long; 
primary bracts like the scape-bracts, much shorter than the divergent to spreading 
branches; spikes short-stipitate, 3-13 cm. long, laxly many-flowered; floral bracts 
broadly ovate, much shorter than the sepals, nerved; flowers sessile, subspreading, 
perfect; sepals elliptic, strongly asymmetric, 6 mm. long; petals slightly exserted; 
style short but distinct. 

Catopsis Morreniana Mez in DC. Monog. Phan. 9: 628. 1896. 
C. Bakeri Mez, Bull. Torrey Club 30: 435. 1903. Figure 75. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 403 




FIG. 75. Catopsis Morreniana. a, Leaf (X Y^). b, Inflorescence and upper 
part of scape ( X H)- c, Sepal ( X 5). d, Petal with three stamens in place ( X 5). 
e, Fruiting branch (X Y^). 



Epiphytic in forests and terrestrial on open ridges, 20-1650 
meters; Pete"n; Alta Verapaz; Chimaltenango. Southern Mexico; 
British Honduras; Honduras; Nicaragua; Costa Rica. 

Plant 2-4 dm. high; leaves 10-18 cm. long, their blades ligulate, apiculate, 
15-25 mm. wide, narrowly white-margined; scape erect, slender, its bracts imbri- 
cate; inflorescence laxly compound, 6-17 cm. long, glabrous, stramineous; primary 
bracts usually shorter than the spikes; spikes stipitate, the staminate many-flow- 
ered, up to 7 cm. long, the perfect or pistillate 4-5 cm. long, densely few-flowered; 
floral bracts ovate, shorter than the sepals; flowers spreading; sepals asymmetric, 
3-6 mm. long. 

Catopsis nitida (Hook.) Griseb. Fl. Brit. W. Ind. 599. 1864. 
Tillandsia nitida Hook. Exot. Fl. pi. 218. 1827. Figure 76. 

Epiphytic in forests, 800-1550 meters; Baja Verapaz (Panjal, 
Tuerckheim 3991}. Honduras; Costa Rica; Panama; Greater An- 
tilles; Guiana. 



404 



FIELDIANA: BOTANY, VOLUME 24 




FIG. 76. Catopsis nitida. a, Flower with floral bract (X 4). b, Corolla 
opened to show stamens (X 4). c, Pistil (X 4). d, Sepal (X 4). e, Leaf-rosette 
and lower part of scape (X %). /, Inflorescence and top of scape (X K)- 

Plant to 45 cm. high; leaves few in an elongate cylindrical rosette, to 35 cm. 
long, sheaths very indistinct, about as long as the blades but scarcely wider, blades 
ligulate, rounded-apiculate, to 4 cm. wide; scape erect or somewhat curved, slender; 
scape-bracts remote, lance-ovate; inflorescence laxly compound with simple or 
rarely divided branches, equaling or exceeding the leaves, 5-20 cm. long; primary 
bracts lanceolate, much shorter than the sterile base of the spike; spikes divergent, 
lax; floral bracts broadly ovate, acute, shorter than the sepals; flowers perfect; 
sepals strongly asymmetric, 5-6 mm. long; petals barely exserted; style lacking. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 405 

Catopsis nutans (Sw.) Griseb. Fl. Brit. W. Ind. 599. 1864. 
Tillandsia nutans Sw. Prodr. 56. 1788. T. vitellina Lk., Kl. & Otto, 
Ic. PI. Rar. 101. 1843. Catopsis fulgens Griseb. in Nachr. Ges. Wiss. 
Gott. 1864: 21. 1865. C. vitellina Baker, Jour. Bot. 25: 176. 1887. 
Gallito quiz (fide Aguilar). Figure 77. 

Epiphytic, 75-1500 meters; Alta Verapaz; Escuintla; Guatemala; 
San Marcos. Greater Antilles; Vera Cruz to Panama, Venezuela 
and Ecuador. 




FIG. 77. Catopsis nutans. a, Flower with bract (X %) b, Petal with 
stamen in position (X %). c, Habit (X M)- 



Plant 14-40 cm. long extended; leaves about 10, up to 24 cm. long, sheaths 
elliptic, blades subtriangular, 25 mm. wide; scape slender, usually decurved, its 
bracts remote; inflorescence simple or rarely few-branched with very short primary 
bracts; spikes lax, up to 2 dm. long; floral bracts shorter than the sepals; flowers 
perfect, erect to spreading; sepals strongly asymmetric, 15 mm. long; petals 2 cm. 
long, bright yellow. 

Catopsis nutans var. robustior L. B. Smith, Contr. Gray 
Herb. 154: 34. 1945. 

Epiphytic, 1300-1500 meters; endemic; Chiquimula; Quezalte- 
nango (type from between Finca Pirineos and Patzulin, Standley 
87157). 

Scape erect or suberect, to 3 mm. thick; scape-bracts less remote or even 
slightly imbricate. 

Catopsis nutans var. stenopetala (Baker) L. B. Smith, No. 
Am. Fl. 19: 193. 1938. Catopsis stenopetala Baker, Jour. Bot. 25: 
176. 1887 (type from Guatemala without further locality, Bernoulli 
& Cario 685). 



406 FIELDIANA: BOTANY, VOLUME 24 

Epiphytic, 600-1200 meters; Santa Rosa; Escuintla; Sacatepe*- 
quez; Huehuetenango. Southern Mexico; El Salvador. 

Flowers functionally dioecious; fruiting pistillate plants indistinguishable from 
the typical form of the species; staminate plants more delicate with ampler inflo- 
rescence, slenderly cylindric calyces and sepals 12 mm. long. 

Catopsis pedicellata L. B. Smith, Contr. Gray Herb. 154: 34. 
1945. 

Epiphytic in pine and oak woods, alt. 500-1500 meters; Chiqui- 
mula (type and only collection from Cerro Tixixi, 3-5 miles north of 
Jocotan, Steyermark 31641). 

Flowering plant to 6 dm. high; leaves many in a cyathiform rosette, to 2 dm. 
long and 45 mm. wide, subelliptic without a distinct sheath, acute or subacute and 
apiculate, narrowly pale-margined, glabrous; scape erect, slender; scape-bracts 
elliptic, acute, erect, all but the uppermost imbricate; inflorescence laxly tripin- 
nate, 19 cm. long; primary bracts ovate, acute, small; branches suberect, lax; floral 
bracts broadly ovate, barely exceeding the pedicels; pedicels slender and quite dis- 
tinct, 1-1.5 mm. long; sepals broadly elliptic, obtuse, subsymmetric, 2.5 mm. long, 
thin; petals 5 mm. long; capsule slender, acute, 12-13 mm. long. 

Catopsis pendula Baker, Handb. Bromel. 155. 1889. 

Terrestrial in swamps, epiphytic in forests, 1300-1575 meters; 
Alta Verapaz; Baja Verapaz. Southern Mexico; Costa Rica. 

Plant up to 1 meter long extended; leaves about 12 in a crateriform rosette, 
3-4 dm. long, sheaths large but scarcely distinct from the blades, elliptic, blades 
triangular, 3-6 cm. wide; scape deflexed, much longer than the leaves, its bracts 
foliaceous, densely imbricate; inflorescence amply tripinnate, 3-5 dm. long or more; 
primary bracts like the scape-bracts, barely exceeding the sterile bases of the weak 
elongate branches; spikes many-flowered, the staminate subdense, 65 mm. long, 
the pistillate lax, up to 25 cm. long; floral bracts broadly ovate, obtuse, much 
shorter than the sepals; flowers functionally dioecious, the staminate 7 mm. long, 
the pistillate up to 12 mm. long; sepals strongly asymmetric, the staminate 5 mm. 
long, the pistillate 5-9 mm. long; petals exceeding the sepals. 

Catopsis sessiliflora (R. & P.) Mez in DC. Monog. Phan. 9: 
625. 1896. Tillandsia sessiliflora R. & P. Fl. Peruv. 3: 42. 1802. 
Catopsis nutans var. erecta Wittm. Bot. Jahrb. 11: 71. 1889. Gallito 
quiz (fide Aguilar). Figure 78. 

Epiphytic, at very low altitudes; Izabal (vicinity of Quirigua, 
75-225 meters, Standley 24546). West Indies; southern Mexico to 
southern Brazil, Colombia and Peru. 

Plant 1-3 dm. high; leaves 4-13 in a slender rosette, 8-20 cm. long, sheaths 
inconspicuous, about as long as the blades but scarcely wider, blades curving out- 




FIG. 78. Catopsis sessiliflora. a, Habit (X Yd. b, Sepal (X 2^). c, Seed 



407 



408 FIELDIANA: BOTANY, VOLUME 24 

ward, ligulate, rounded and apiculate, 12-25 mm. wide; scape erect, slender; scape- 
bracts erect, remote; inflorescence simple or compound from a few spikes, lax, to 

11 cm. long; primary bracts much shorter than the sterile bases of the spikes; spikes 
divergent, laxly flowered; floral bracts broadly ovate, obtuse, much shorter than 
the sepals; flowers suberect, perfect; sepals asymmetric, suborbicular, 7-8 mm. 
long, thin; petals barely exserted, white; style very short; capsule ovoid, distinctly 
short-beaked, 12 mm. long. 

Catopsis subulata L. B. Smith, Contr. Gray Herb. 114: 5. 1936. 

Epiphytic in pine-oak forests, 1300-2000 meters; Baja Verapaz; 
Huehuetenango. Southern Mexico; Honduras. 

Plant 3-6 dm. high; leaves many in an ellipsoid pseudobulb, 15-30 cm. long, 
sheaths about twice as long as the blades, ample, blades narrowly triangular, acu- 
minate, 1 cm. wide, involute and contorted; scape erect, exceeding the leaves, its 
bracts imbricate, lanceolate, involute, recurving toward apex; inflorescence com- 
pound, rather dense, 6-21 cm. long; primary bracts much shorter than the branches, 
broadly ovate with an acuminate recurved apex; branches simple or the lowest 
divided; spikes cylindric, dense, 2-7 cm. long; floral bracts broadly ovate, acute, 
much shorter than the sepals, thin; flowers functionally dioecious, suberect; sepals 
strongly asymmetric, 9 mm. long; petals included, white. 

Catopsis triticea L. B. Smith, Contr. Gray Herb. 127: 19. 1939. 

Epiphytic in wet forests, 1360-1450 meters; Alta Verapaz. 
Chiapas. 

Plant 2-3 dm. high; leaves 8-12 cm. long, narrowly white-margined, sheaths 
large, elliptic, blades narrowly triangular, acuminate, about 15 mm. wide; scape 
erect, slender, exceeding the leaves; scape-bracts imbricate, broadly ovate; stami- 
nate inflorescence 6.5 cm. long, sparsely branched at base; primary bracts much 
shorter than the spikes; spikes dense; floral bracts ovate, acute, straight, about 
equaling the sepals, nerved; flowers 10 mm. long; sepals 9 mm. long, asymmetric; 
petals exserted; pistillate inflorescence simple, dense, 6-8 cm. long; floral bracts 

12 mm. long, exceeding the sepals; flowers 13 mm. long; sepals 10 mm. long, asym- 
metric; style 1 mm. long. 

Catopsis Wawranea Mez in DC. Monog. Phan. 9: 626. 1896. 
Figure 79. 

Epiphytic in high forest; southern Mexico; British Honduras; 
Costa Rica; probably extending into Guatemala. 

Plant up to 35 cm. high; leaves few, 2 dm. long, blades ligulate, apiculate, 4 cm. 
wide; scape suberect to decurved; scape-bracts remote; inflorescence few-branched 
or simple; primary bracts short; floral bracts narrow, thin, shorter than the sepals; 
flowers suberect, dioecious; sepals subsymmetric, thin, the staminate lanceolate, 
acute, 15 mm. long, the pistillate broadly elliptic, subobtuse, 18 mm. long. 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 409 




FIG. 79. Catopsis Wawranea. a, Inflorescence and upper part of scape with 
apex of leaves (X V^}. b, Pistillate sepal (XI). 



GREIGIA Regel 

Coarse terrestrial or saxicolous herbs, usually stemless or short-caulescent; 
leaves many in a dense rosette, blades linear; scape none; inflorescence central or 
lateral, compound or simple, dense; floral bracts evident; flowers perfect; sepals 
free or short-connate at base, petals connate by their margins; stamens usually 
included, filaments connate with the petals, pollen globose, smooth; ovary triangu- 
lar, epigynous tube very short or none; ovules numerous; fruit fleshy. 

Eighteen species from Mexico to Chile and Juan Fernandez. 



Greigia Steyermarkii L. B. Smith, Contr. Gray Herb. 154: 35. 
1945. Figure 80. 

Cloud forest, 2100-2400 meters; endemic; Zacapa (type from 
mountain above Rio Repollal, Steyermark 1^2553} . Mexico. 

Stem stout, 1.5-2.4 meters long; leaves 1 meter or longer, spreading or ascend- 
ing, sheaths narrowly triangular-ovate, dark castaneous, covered with a whitish 
membrane outside, subglabrous inside, subdensely serrate toward apex with stout 
dark spines up to 4 mm. long, blades long-acuminate, slightly constricted and 
channeled toward base, to 3 cm. wide, glabrous above, whitish-lepidote beneath, 
armed with pale spines to 1.5 mm. long; inflorescence dense, few-flowered; outer 
bracts triangular-ovate, acuminate, pungent, dark castaneous, whitish-lepidote, 
coarsely serrate; floral bracts lanceolate, acuminate, entire, carinate, pungent, 26 
mm. long, exceeded by the sepals, whitish-lepidote, castaneous toward apex; flowers 
4 cm. long; sepals like the floral bracts, 22 mm. long; petals 30-32 mm. long, white; 
stamens and style included. 




410 



STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 411 

GUZMANIA Ruiz & Pavon 

Stemless or rarely long-caulescent herbs, mostly epiphytic; leaves entire, the 
sheaths conspicuous; inflorescence simple or compound, the spikes always poly- 
stichous-flowered; flowers perfect; sepals usually somewhat connate; petals con- 
nate or closely agglutinated, naked, yellow or white; stamens usually included, the 
filaments more or less fused to the petals; ovary wholly superior; ovules many; 
seeds with a long basal straight usually brownish coma. 

One hundred and ten species in tropical and subtropical America. 

Inflorescence simple. 

Inflorescence cyathiform, the outer bracts much enlarged, red. 
Leaves not over 25 mm. wide; inner bracts nearly similar to the outer. 

G. minor 

Leaves much wider; inner bracts much reduced G. lingulata 

Inflorescence densely fusiform G. nicaraguensis 

Inflorescence compound, with lax elongate spikes. 

G. Scherzeriana var. guatemalensis 

Guzmania lingulata (L.) Mez in DC.